Ranitomeya Bauer 1988

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), pp. 1-120 : 39-40

Ranitomeya Bauer 1988

Account authors: J.L. Brown, E. Twomey

Figs. 3 , 4 , 9–45 , Tables 2–7

Type species. Dendrobates reticulatus Boulenger, 1884 “1883”

Proposed sister group. Andinobates gen. nov.

Definition and diagnosis. Unambiguous synapomorphies include: 5 nuclear and 16 mitochondrial synapomorphies (based upon the dataset used in this study, Fig. 3 , Tables 2 and 3); distinctive pale reticulation on limbs and venter present ( Fig. 2 c-i and Fig. 2 h-i). Secondary losses of this pale limb reticulation appear to have occurred in R. yavaricola and R. summersi , some morphs of R. fantastica , R. imitator and R. sirensis . Other features include: adult SVL less than 21 mm; adults typically brightly colored, often with bright yellow, red, or green dorsal coloration (either uniform, spotted, or striped); dorsolateral stripe, if present, extending to top of thigh (vs. not reaching thigh in Andinobates ), ventrolateral stripe and oblique lateral stripe present or absent; Distinctive, bright coloration on throat present (usually yellow, orange or red); dorsal skin texture nearly smooth to weakly granular; head narrower than body; teeth absent; vocal slits present in males; finger I greatly reduced and shorter than finger II; finger discs II–IV greatly expanded; in adults disc on finger III at least two times wider than distal end of adjacent phalanx; thenar tubercle conspicuous (commonly vestigial, occasionally absent); toe discs III–V moderately expanded; toe webbing absent; median lingual process absent; larval vent tube dextral; larval oral disc emarginated; larvae without medial gap in papillae on posterior labium (known in all species expect R. cyanovittata and R. ventrimaculata , Table 4); scansorial; adults use arboreal phytotelmata for reproduction and deposit eggs away from or at edge of water in phytotelmata (Table 6); tadpoles deposited individually, typically by male; small clutches (2– 6 eggs, Table 6); vertebrae 2 and 3 unfused (known in R. amazonica , R. toraro sp. nov., R. imitator , R. variabilis , R. vanzolinii , R. fantastica , R. reticulata and R. sirensis ).

Distribution. This genus occurs within Amazonian rainforests of Brazil (States: Acre, Amapá, Amazonas, Pará, Rondônia), Bolivia (Department: Pando), Colombia (Departments: Amazonas, Caquetá, Putumayo (tentative), Vaupés), Ecuador (Provinces: Morona-Santiago, Napo, Orellana, Pastaza, Sucumbíos), French Guiana (Arrondissements: Cayenne, Saint-Laurent-du-Maroni), Guyana (District: Potaro-Siparuni) and Peru (Departments 1: Amazonas, Cusco, Huánuco, Junín, Loreto, Madre de Dios, Pasco, San Martín, Ucayali). Species within this genus occur between sea level and 1600 m, Fig. 9 .

Species included (16). Dendrobates amazonicus Schulte 1999 ; Ranitomeya benedicta Brown, Twomey, Pepper & Sanchez-Rodriguez 2008 ; Ranitomeya cyanovittata Perez-Peña, Chavez, Twomey & Brown 2010 ; Ranitomeya defleri Twomey & Brown 2009 ; Dendrobates fantasticus Boulenger 1884 “1883”; Dendrobates flavovittatus Schulte 1999 ; Dendrobates imitator Schulte 1986 with its junior synonyms Dendrobates imitator intermedius Schulte 1999 and Dendrobates imitator yurimaguensis Schulte 1999 ( Vences & Lötters 2000, Lötters et al. 2003

1. Since 2009, Peru officially reclassified ‘Departmentos’ as ‘Regiones’. At the time of writing, however, ‘region’ is infrequently used in scientific literature. Here we use Departments as a synonym of Regions.

and this paper); Dendrobates reticulatus Boulenger 1884 “1883” with its junior synonym Dendrobates tinctorius igneus Melin 1941 ; Dendrobates sirensis Aichinger 1991 with its junior synonyms Dendrobates biolat Morales 1992 and Dendrobates lamasi Morales 1992 (this paper); R. summersi Brown, Twomey, Pepper & Sanchez-Rodriguez 2008 ; Ranitomeya toraro sp. nov. (this paper); Dendrobates uakarii Brown, Schulte & Summers 2006 ; Dendrobates vanzolinii Myers 1982 ; Dendrobates variabilis Zimmermann & Zimmermann 1988 ; Dendrobates ventrimaculatus Shreve 1935 with its junior synonym Dendrobates duellmani Schulte 1999 (this paper); Ranitomeya yavaricola Perez-Peña, Chavez, Twomey & Brown 2010 .

Remarks. Our definition of Ranitomeya is essentially equal to the definition of Caldwell & Myers (1990) ventrimaculatus group. The genus apparently diverged from Andinobates approximately 14 mya during the mid-Miocene ( Santos et al. 2009).

Figures

FIGURE 2. Illustrated guide to morphological terminology. A. Finger and hand morphology: i. Finger I (far left) <Finger II, thenar tubercle (= inner metacarpal tubercle) present (depicted by arrow), and greatly expanded finger discs in Fingers II-IV. Inset depicts Finger I and a thenar tubercle which is clearly visible. Note that in some Ranitomeya this is trait reduced and difficult to view (as in main picture) (Ranitomeya variabilis pictured, inset of R. benedicta). ii. Finger I ≈ Finger II, thenar tubercle absent. (Adelphobates quinquevittatus pictured) iii. Weakly expanded finger discs in Fingers II-IV (Excidobates captivus pictured). B. Stripes: i. Middorsal (follows vertebral column), dorsolateral (extends from eye to either upper thigh, as pictured, or to vent), ventrolateral (running from groin to axilla) and labial stripe (stripe that extends from shoulder around upper lip)(R. sirensis pictured). ii.. Oblique lateral stripe (extends from groin to eye, as in picture stripe is incomplete anteriorly). Unlabeled arrow depicts a dorsolateral stripe that does not reach thigh, a characteristic of certain species of Andinobates (type ‘A’ in Grant et al. 2006). (Andinobates claudiae pictured). C. Limb patterns: i. Distinct limb reticulation/spotting (characteristic of most species of Ranitomeya) (R. variabilis pictured). ii. Wavy stripes (not classified as distinct limb reticulation) (R. summersi pictured). iii. Patternless. Typical of most Andinobates species (R. sirensis pictured). D. Diagnostic head patterns: i. Large black “oval” on head (R. imitator pictured). ii. Large black “pentagon” or “five-point star” on head (R. summersi pictured). iii. Black band across head entirely covering eyes (known only in a single population of this species near the Pongo de Manseriche, Peru) (R. fantastica pictured). E. Nose spots. i. Two nose spots (R. imitator pictured). ii. Single nose spot. (R. variabilis pictured). iii. Frontward-turned “U” on the tip of snout. (R. toraro pictured). F. Geographical distribution. West: distribution within Andes, west of Andes, or in Central America. East: distribution east of Andes (including Guiana Shield) or in east-Andean versant. G. Dorsal patterns: i.“Y-shape”. Space between stripes create black pattern which forms a black Y on the back. (R. variabilis pictured). ii. Merging of the obliquelateral and dorsolateral stripes (R. variabilis pictured). iii. Broken dorsolateral stripes (R. flavovittata pictured). iv. Spotting (R. imitator pictured). H. Key ventral characters: i. Distinctive throat coloration and ventral reticulation (also shown in H-ii & H-iii) (R. reticulata pictured). ii. Belly patch (R. sirensis pictured). iii. Gular spots (single or paired dark spots at corner of mouth) (R. amazonica pictured). iv. Marbled pattern (not classified as reticulation) (Andinobates virolinensis pictured).

FIGURE 3. A consensus Bayesian phylogeny based on 1011 base pairs of aligned mitochondrial DNA sequences of the 12S (12s rRNA), 16S (16s rRNA) and cytb (cytochrome-b gene) regions. Thickened branches represent nodes with posterior probabilities 90 and greater, other values are shown on nodes. Taxon labels depict current specific epithet, number in tree, the epithet being used prior to this revision (contained in parentheses), and the collection locality. A. Top segment. B. Middle segment. C. Bottom segment of phylogeny.

FIGURE 4. Putative species tree for Andinobates, Excidobates, and Ranitomeya. Placement of species where molecular data were lacking (A. altobueyensis, A. viridis, A. abditus, A. daleswansoni and R. opisthomelas) was based on morphology. Andinobates altobueyensis and A. viridis were placed as sister taxa due to the absence of dark pigmentation on dorsal body and limbs and overall similar dorsal coloration and patterning. These species were placed as sister to A. fulguritus (sequenced) on the basis of similar dorsal coloration (bright green to greenish-yellow). Andinobates opisthomelas was placed in the bombetes group in a polytomy with A. bombetes and A. virolinensis (both sequenced) due to their similar advertisement calls and morphology, particularly their red dorsal pattern and marbled venter. Andinobates daleswansoni was placed as sister to A. dorisswansonae due to the absence of a well-defined first toe in both species. Andinobates abditus was placed in the bombetes group based on a larval synapomorphy which appears to be diagnostic of that group (wide medial gap in the papillae on the posterior labium). However, A. abditus was placed as the sister species to all other members of the bombetes group due to the absence of bright dorsal coloration and isolated geographic distribution. Andinobates abditus is currently the only species of its genus known to occur in the east-Andean versant, thus its placement remains speculative until molecular data become available. Photo credits: Thomas Ostrowski, Karl-Heinz Jungfer, Victor Luna-Mora, Giovanni Chaves-Portilla.

FIGURE 9. Known elevation distributions of Ranitomeya. Dotted line is mean for all samples. Dark boxes display the total elevation range of each species, within each contains a corresponding box plot.

FIGURE 10. Ranitomeya Plate 1. defleri group: A–B: Ranitomeya defleri (all from Vaupés, Colombia); A: Holotype at MCZ (Ω); B: near Estación Biológica Caparú (1 Φ). C–M: Ranitomeya toraro from Brazil; C–D: Careiro da Varzea, Amazonas (A. P. Lima); E: Humaitá, Amazonas (P. I. Simoes); F–G: Cachoiera do Jirau, Rondônia (W. Hödl); H: near Boca do Acre, Amazonas (PRMS and MBS, Ω); I: Host plant of R. toraro: Phenakospermum guyanense near Boca do Acre, Acre (MBS); J: near Boca do Acre, Amazonas (PRMS and MBS); K: Habitat of R. toraro near Boca do Acre, Acre, inset: Aechmea sp. used for tadpole deposition (MBS); L: near Boca do Acre, Amazonas (PRMS and MBS); M: Rio Ituxi, Amazonas (JPC, Ω). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny).

FIGURE 11. Ranitomeya Plate 2. defleri group: A–H: Ranitomeya toraro (all from Brazil); A-B: Rio Branco, Acre (T. Grant); C-F: Upper Jurua, Acre (unknown). From Colombia: G: Leticia, Amazonas (Jose Manuel Padial, Ω); H: Axil of Aechmea sp. with two R. toraro embryos, near Boca do Acre, Amazonas, BZ (MBS). I: Adelphobates quinquevittatus, near Boca do Acre, Amazonas, BZ (PRMS); J: R. uakarii near Porto Walter, Acre, BZ (JPC, Ω); K: Tadpole of R. toraro (MBS). (Ω = population sampled in phylogeny).

FIGURE 12. Ranitomeya toraro sp. nov. type series. All specimens from two localities in Brazil: Amazonas state, municipality of Castanho, at km 12 on road to Autazes (ca. 40 km south of Manaus) or Scheffer Madeireira on Rio Ituxi, ca. 170 km southwest of Lábrea (labeled with ‡). Top row, from the collections of MPEG (L-R): 13839, 13838 (holotype), 13841, 13840, 13842 and 13037(‡). Bottom row, from the collections of OMNH (L-R): 37441, 37440, 36666 (‡), 37442, 37439, 36667(‡), 37438. Black bar = 20 mm (5 mm increments). Sequenced individuals (number in phylogeny): OMNH 36666 (7), OMNH 37440 (5), OMNH 36667 (6), MPEG 13841 (4)

FIGURE 13. Known distribution of the defleri group. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 14. Advertisement calls of Ranitomeya species in the variabilis group and defleri group. A. Ranitomeya amazonica from 23 km S Iquitos, Loreto, Peru (type locality), recorded at 26° C; B. Ranitomeya amazonica from French Guiana, unknown temperature (call courtesy Erik Poelman); C. Ranitomeya variabilis from Cainarachi valley, San Martín, Peru, recorded at 22° C. D. Ranitomeya variabilis from Cerro Yupatí, Amazonas, Colombia, recorded at 27° C; E. Ranitomeya variabilis from Saposoa, San Martín, Peru, recorded at 24.5 C; F. Ranitomeya defleri from Rio Apaporis, Vaupés, Colombia, recorded at 26° C.

FIGURE 15. Ranitomeya Plate 3. reticulata group: A–D: Ranitomeya benedicta (all from Peru); A–B: Shucushuyacu, Loreto (1Φ); C-D: Pampa Hermosa, Loreto. E–L: Ranitomeya fantastica (all from Peru); E: Yurimaguas, Loreto; F: near Yumbatos, San Martin; G: Pongo de Cainarachi, San Martin (Ω); H: Cainarachi Valley, San Martin (Ω); I: San Antonio, San Martin (KS); J: Tarapoto, San Martin (Ω); K: Santa María de Nieva, Loreto (K.-H. Jungfer, 1Φ); L: Lower Huallaga Canyon, San Martin (Ω). M & N: Ranitomeya summersi (all from San Martin, Peru); M: Chazuta (3Φ); N: Sauce (Ω). O–R: Ranitomeya reticulata (all from Loreto, Peru); O-P: Iquitos (Ω); Q: Puerto Almendras (PPP); R: Upper Rio Itaya (PPP). (nΦ= number of individual in phylogeny, Ω = population sampled in phylogeny).

FIGURE 16. Ranitomeya Plate 4. reticulata group: A–G: Ranitomeya reticulata (all from Loreto, Peru); A: Lower Rio Itaya (Ω); B–D: Upper Itaya Drainage; E-F: Rio Sucusari (KS); G: 30 km west of Pevas (MSR). H–M: Ranitomeya ventrimaculata: H: Kapawi, Pastaza, EC (L. Coloma, 7Φ); I: Upper Curaray Drainage, Loreto, PE (PPP); J: Yasuní, Orellana, EC (A. Blasco Z., Ω); K: central Rio Nanay (M. Callegari, 6Φ); L: Holotype at MCZ, Sarayacu, Pastaza, Ecuador; M: Yasuní, Orellana, EC (J. Yeager, Ω); N: Yasuní, Orellana, EC (S. Ron, Ω). O & P: Ranitomeya uakarii (all from Loreto, Peru): O: Tamshiyacu village (8Φ) P: Quebrada Blanco (12Φ). (nΦ = number of individual in phylogeny;Ω = population sampled in phylogeny).

FIGURE 17. Ranitomeya Plate 5. reticulata group: A–I: Ranitomeya uakarii: A: Central Rio Yavari, Loreto, Peru (PPP); B: Quebrada Blanco, Loreto, Peru (PPP, Ω); C: Rio Boncuya, Loreto, Peru (G. Gagliardi); D: Tournavista, Huánuco (A. Toebe); E: Porto Walter, Acre, Brazil (JPC); F: Rio Los Amigos, Madre de Dios, Peru (RVM); G: R. uakarii sp. aff. Caquetá, Colombia (J. M. Rengifo); H: R. uakarii sp. aff. Iwokrama, Guyana (unknown photographer); I: Guzmania bromeliad with R. uakarii embryos near Tamshiyacu village, Loreto, Peru. (Ω = population sampled in phylogeny).

FIGURE 18. Advertisement calls of Ranitomeya species in the fantastica group. A. Ranitomeya reticulata from Iquitos, Loreto, Peru, recorded at 29° C; B. Ranitomeya ventrimaculata, unknown locality or temperature; C. Ranitomeya fantastica from Cainarachi valley, San Martín, Peru, recorded at 24° C; D. Ranitomeya summersi from Sauce, San Martín, Peru, recorded in captivity at 24.5° C; E. Ranitomeya benedicta from Shucushuyacu, Loreto, Peru, recorded in captivity at 26.5° C; F. Ranitomeya uakarii from Rio Tahuayo, Loreto, Peru, recorded in captivity at 26° C.

FIGURE 19. Known distribution of Ranitomeya benedicta and R. summersi. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 20. Known distribution of Ranitomeya fantastica. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 21. Known distribution of Ranitomeya reticulata and R. ventrimaculata. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 22. Known distribution of Ranitomeya uakarii. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 23. Ranitomeya Plate 6. vanzolinii group: A & B: Ranitomeya cyanovittata: Sierra del Divisor, Ucayali, Peru (G. Knell and D. Vasquez, 1:Ω,2: 1Φ). C & D: Ranitomeya yavaricola (all from Loreto, Peru): C: Rio Blanco (G. Knell); D: Lago Preto (PPP, Ω). E– I: Ranitomeya flavovittata (all from Quebrada Blanco, Loreto, Peru (Photo credits: JLB, ET and PPP, Ω). J–K: Ranitomeya vanzolinii Atalaya, Ucayali, Peru (J. Yeager). L–V: Ranitomeya imitator (All from San Martin, Peru): L–O: Upper Canarachi Valley (‡); P– Q: Tarapoto (‡); R: Shapaja (‡); S: Chumia (‡) and T–V: Chazuta (Ω). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny, ‡ = genetically sampled, but not included in our phylogeny).

FIGURE 24. Ranitomeya Plate 7. vanzolinii group: A–Y: Ranitomeya imitator (all from San Martin, Peru unless noted): A-B: Chazuta (Ω); C: Central Huallaga Canyon (‡); D-H: Callanayacu (‡); I-J: Lower Huallaga Canyon (‡); K-Q: Pongo de Cainarachi (Ω); R- S: Balsapuerto, Loreto (‡); T–V: Varadero, Loreto (‡) and W–Y: Curiyacu (‡). (Ω = population sampled in phylogeny, ‡ = genetically sampled, but not included in our phylogeny).

FIGURE 25. Ranitomeya Plate 8. vanzolinii group: A–I: Ranitomeya imitator Curiyacu, San Martin, Peru (‡). J–T: Ranitomeya sirensis (all from Peru unless noted): J-L: CICRA Station, Madre de Dios (Rio Los Amigos, Ω); M: near Rio Branco, Acre, Brazil (PRMS); N & O: Central Rio Urubamba, Cusco (G. Chavez); P & Q: Tingo Maria, Huánuco (ET, Ω); R: Bamboo forest, R. sirensis often uses the phytotelmata within bamboo for tadpole deposition, Tingo Maria, Huánuco (ET); S: Aguaytía, Ucayali; T: Codo del Pozuzo, Huánuco (20Φ). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny, ‡ = genetically sampled, but not included in our phylogeny).

FIGURE 26. Ranitomeya Plate 9. vanzolinii group: A–P: Ranitomeya sirensis (all from Peru): A–G: Puerto Inca, Huánuco (JLB and ET, Ω); H & I: Breeding pair of R. sirensis found in the type locality between the lowland and highland populations, Cordillera El Sira, Huánuco (MSR); J & K: Rio Pachitea, Huánuco (J. Stenicka); L–N: Cordillera El Sira, Huánuco (B. Wilson and JLB, 10-11Φ); O: Rio Pachitea, Huánuco (J. Stenicka, 17Φ); P: Yanayacu Maquia, Ucayali. (nΦ = number of individual in phylogeny Ω = population sampled in phylogeny).

FIGURE 27. Ranitomeya Plate 10. vanzolinii group: A–F: Ranitomeya sirensis (all from Peru): A–D: near Contamana, Loreto (JLB and G. Gagliardi, 1-3Φ); E: uncertain locality, likely Iscozacin, Junin (7 Φ); F: Estación Biológica Paujil, Junin (L. Schulte). (nΦ = number of individual in phylogeny).

FIGURE 28. Advertisement calls of Ranitomeya species in the vanzolinii group. A. Ranitomeya vanzolinii from Pongo de Mainique, Cuzco, Peru, recorded in captivity at 26° C; B. Ranitomeya sirensis from Ishanga near Tocache, San Martín, Peru, recorded at 25.5° C; C. Ranitomeya imitator from Varadero, Loreto, Peru, recorded at 23.5° C; D. Ranitomeya imitator from Cainarachi valley, San Martín, Peru, recorded at 22 C; E. Ranitomeya flavovittata from Rio Tahuayo, Loreto, Peru, recorded in captivity at 25° C; F. Ranitomeya yavaricola from Lago Preto, Loreto, Peru, recorded at 24° C (call courtesy Pedro Pérez-Peña).

FIGURE 29. Known distribution of Ranitomeya flavovittata, R. yavaricola and R. cyanonvittata. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 30. Known distribution of Ranitomeya imitator. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 31. Known distribution of Ranitomeya sirensis and R. vanzolinii. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 32. Ranitomeya Plate 11. variabilis group. A–P: Ranitomeya variabilis (Highland morph): A: Chazuta, San Martin, Peru; B–G: Upper Cainarachi Valley, San Martin, Peru (Ω); H & I: Borja, Loreto, Peru (24, 46Φ); J: Parque Nacional Ichigkat Muja, Amazonas (D. Rodriquez-Mercado); K: Saposoa, San Martin, Peru (Ω); L: Comparision between Saposoa and upper Cainarachi Valley populations (respectively); M: Xanthosoma sp. that was being used by R. variabilis for tadpole and egg deposition near Saposoa, San Martin, Peru; N: Saposoa, San Martin, Peru (Ω); O: Tocache, San Martin, Peru (C. Torres); P: Macas, Morona Santiago, Ecuador (J. Verkade, Ω). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny).

FIGURE 33. Ranitomeya Plate 12. variabilis group: A–P: Ranitomeya variabilis (Lowland morph): A: Contamana, Loreto, Peru (42Φ); B: Shamboyacu, San Martin, Peru (41Φ); C: Lower Huallaga Canyon, San Martin, Peru (‡); D: Callanayacu, San Martin, Peru(‡); E: Barranquita, San Martin, Peru (‡); F: Pongo de Cainarachi, San Martin, Peru (7,8,11Φ); G & H: Bonilla, San Martin, Peru (1,3, 9 Φ); I & J: Varadero, Loreto, Peru (27Φ); K: Quebrada Blanco, Loreto, Peru (Ω); L: Rio Boncuya, Loreto, Peru (G. Gagliardi); M: Upper Rio Nanay, Loreto, Peru; N: Middle Rio Nanay, Loreto, Peru; O & P: Archidona, Napo, Ecuador (EHP and J. Verkade). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny).

FIGURE 34. Ranitomeya Plate 13. variabilis group: A–K: Ranitomeya variabilis: A: Archidona, Napo, Ecuador (J. Verkade); B– D: Macuma, Morona-Santiago, Ecuador (J. Verkade and EHP); E &F: Puyo, Pastaza, Ecuador (j. Verkade) G: Yupati, Vaupés, Colombia (14 Φ); H: Embryos, Archidona, Napo, Ecuador (J. Verkade); I: Breeding pair of R. variabilis, Archidona, Napo, Ecuador (J. Verkade); J & K: Yupati, Vaupés, Colombia (14-16 Φ). L–Q: Ranitomeya amazonica: L: Km 41 Iquitos, Loreto, Peru (ET, 26Φ); M: Km 31 Iquitos, Loreto, Peru (PPP); N & O; Km 26 Iquitos, Loreto, Peru (ET, 25, 27Φ); P: Upper Rio Mazan-Pintuyacu, Loreto, Peru (J. J. Lopez-Rojas); Q: Lower Rio Mazan, Loreto, Peru (J. J. Lopez-Rojas). (nΦ = number of individual in phylogeny).

FIGURE 35. Ranitomeya Plate 14. variabilis group: A–Q: Ranitomeya amazonica: A–C: Iquitos, Loreto, Peru; D–K: ‘Arena Blanca’, Loreto, Peru (16–18, 20–24Φ); L: 30 km west of Pevas, Loreto Peru; M: Rio Sucusari, Loreto, Peru (B. Pieper); N & O: Nouragues French Guiana (EHP); P: French Guiana (B.P. Noonan, Ω); Q: Estação Científica Ferreira Penna-Caxiuanã, Para, Brazil.(nΦ = number of individual in phylogeny, Ω=population sampled in phylogeny).

FIGURE 36. Known distribution of Ranitomeya amazonica. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 37. Known distribution of Ranitomeya variabilis. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya).

FIGURE 38. Type series of Dendrobates rubrocephalus Schulte 1999. A. Holotype (BD 5H, MHNJP 408) and B. Paratype (BD 6P, MHNJP 408).

FIGURE 39. A consensus Bayesian phylogeny based on 2262 base pairs of nuclear DNA from five genes (Rag1, Rhodopsin, Histone H3, 28s, 7th in Absentia). Thickened branches represent nodes with posterior probabilities higher than 85 (those which are not labeled are 100). In addition, some branches of interest with posterior probabilities lower than 85 are labeled.

FIGURE 40. Consensus of 820 most parsimonious trees for 2622 base pairs of five nuclear genes (Rag1, Rhodopsin, Histone H3, 28s, 7th in Absentia). Thickened branches represent nodes with a consensus percentage of 100%.

FIGURE 41. Phylogenetic relationships of genera. Left: Relationships between genera of Grant et al. 2006 based on ca. 6100 bp of nDNA and mtDNA, optimality criterion: maximum parsimony, numbers on branches depict Bremer support values. Center: Relationships between genera of Twomey & Brown 2008 based on 2124 bp of nDNA and mtDNA (Bayesian analysis) and Santos et al.2009 based on 2895 bp of nDNA and mtDNA (Bayesian and maximum likelihood analyses). Number on branches depict: posterior probabilities from Twomey & Brown 2008 / bootstrap values from Santos et al. 2009 / posterior probabilities from Santos et al. 2009. Right: Relationships between genera from this study based on 1011 bp of mtDNA, Bayesian analysis, number on branches depict posterior probabilities. If genus was excluded, focal study lacked corresponding taxa.

FIGURE 42. Ranitomeya tadpole morphology. Major features of: A. Oral disc morphology, B. Body coloration, C. Body morphology.

FIGURE 43. Potential examples of mimicry in Ranitomeya. A. Ranitomeya uakarii and R. toraro from western Brazil. B. Ranitomeya amazonica and R. reticulata from undisclosed locality in Loreto, Peru. C. Ranitomeya imitator, R. variabilis, R. fantastica from Varadero, Loreto, Peru. In this population, individuals of R. imitator display a cline of morphologies, usually closely resembling local morphs of either R. variabilis or R. fantastica. D. Ranitomeya imitator, R. variabilis and R. fantastica from Pongo de Cainarachi, San Martin, Peru. In this population (and surrounding areas), individuals of R. imitator and R. fantastica (nearby populations possess considerably more orange on the head and dorsum and lack the appearance of 2 black stripes down the dorsum) appear similar to widespread lowland morph of R. variabilis. E. Ranitomeya imitator and R. variabilis from Cainarachi Valley, San Martin, Peru. F. Ranitomeya imitator and R. summersi from Sauce, San Martin, Peru. See Symula et al. 2001 for analyses of D, E and F as cases of Müllerian mimicry.