Ranitomeya reticulata Boulenger, 1884

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 : 52-53

Ranitomeya reticulata Boulenger, 1884 “1883”

Account authors: J.L. Brown, E. Twomey, S. Lötters, P. Perez-Peña

Figs. 3 , 4 , 9 , 15 (o – r), 16 (a – g), 18, 21

Tables 1, 4 – 6

Dendrobates reticulatus Boulenger, 1884 “1883”: p. 635, Plate 57, drawing 2 [NHML 1947.2.15.5 – 12 (eight syntypes) collected by Paul Hahnel from “Yurimaguas, Huallaga River, Peru ”]; – Myers & Daly 1980: p. 20; Zimmermann & Zimmermann 1984: p. p. 35, 1985; Almendariz 1987: p. 77; Hermann 1988: p. 78; Wiltenmuth & Nishikawa 1994: p. 57; Divossen 1999: p. 58, 2000: p. 20; Rodriguez & Duellman 1994: p. 16; Lötters et al. 2003: p. 1909; Christmann 2004: p. 6, Figs. on p. 87, 92, 96; Santos et al. 2009, by implication

Dendrobates tinctorius igneus Melin, 1941: p. 66 , Fig. 37A–B [MHNG 19.1.1925,

20.1.1925 (two syntypes) from " Rio Itaya (near Iquitos), Perú ", collected in 1925 ]

Dendrobates quinquevittatus (non Steindachner 1864) – Silverstone 1975 (partim): p. 33, Fig. 14 A–C ; Lescure & Bechter 1982: p. 26

Ranitomeya reticulata — Bauer 1988: p. 1; Grant et al. 2006: p. 171; Lötters et al. 2007: p. 489, Figs. 619, 620; von May et al. 2008a: p. 394, Appendix 1; Werner et al. 2010, 2011: p. 16, Figs. 1–3

Ranitomeya ignea — Grant et al. 2006: p. 171

Dendrobates igneus — Santos et al. 2009, by implication

Background information. Ranitomeya reticulata shares, with R. fantastica , the distinction of being the first species of Ranitomeya to be described ( Boulenger 1884 “1883”). This species is also the type species of the genus Ranitomeya ( Bauer 1988; for discussion see Grant et al. 2006). In the last 30 years, little controversy has surrounded the validity of this species. For a comparison of R. reticulata to R. ventrimaculata , see the R. ventrimaculata account.

Tadpole. The description is based on a single live tadpole from Yanamono, Loreto, Peru. Mouthparts were verified in another tadpole from the same locality. The tadpole was feeding on detritus .

Tadpole in stage 30; no external gills. Body ovoid in dorsal view, wider near vent. Total length 16.6; body length 9.8; tail length 6.8, tail length 41% of total length. Body width 5.1; body depth 4.0, 78% of body width. Eye well-developed; naris small; distance from naris to anterior edge of eye 0.7. Eye positioned dorsally on head, directed dorsolaterally. Spiracle well developed; vent tube dextral.

Tip of tail bluntly rounded. Tail muscle height at base of tail 2.5; tail muscle width at base of tail 2.3; maximum tail height 4.1. Dorsal fin same height as ventral fin.

Oral disc ventral, emarginate; transverse width 2.4, 14% of body width. Single row of small papillae present laterally and ventrally; wide dorsal gap where papillae absent. LTRF 2(2)/3(1) with A-1 developed on upper labium, A-2 with wide medial gap (30% of total width); P-1 on lower labium with narrow medial gap; P-2 equal in width to P-1; P-3 80% width of P-1.

In life, head and body gray; mouthparts visible from above. Abdomen mostly transparent; intestinal coils black, heart visible. Tail musculature uniform gray, dorsal and ventral fins opaque gray.

Natural history. Some authors have observed this species engaging in biparental care and maternal egg provisioning in captivity ( Zimmermann & Zimmermann 1984; Christmann 2004), such as has been observed in the wild in R. imitator and R. vanzolinii ( Caldwell 1997, Caldwell & de Oliveira 1999, Brown et al. 2010). These behaviors have never been confirmed in the field in R. reticulata , despite considerable attention by researchers ( Divossen 1999, 2000; Werner et al. 2011). In these studies, the authors observed a species with male-only parental care and a polygamous mating system. Lastly, other members of the reticulata group (where parental care behaviour is known) demonstrate male-only parental care and biparental care is limited to some members of the vanzolinii group ( Summers & McKeon 2004; Lötters et al. 2007). The phylogenetic placement of this species, sister to other male parental care species, cannot exclude the possibility of the independent evolution of this trait; however, it is not likely, given that biparental care/maternal provisioning is only known to have evolved twice in the family Dendrobatidae ( Summers & McKeon 2004) .

Taxonomic notes. In 1941, Melin described Dendrobates tinctorius igneus . Little information is published on this putative taxon, possibly because of a lack of credence in its validity (see Silverstone 1975). Schulte (1999) regarded it as a junior synonym of R. reticulata , and stated that it was a morph that maintained juvenile coloration. Grant et al. (2006) elevated this subspecies to specific status as Ranitomeya ignea , without comment or justification. After examining the holotype, Lötters & Vences (2000) suggested this species may be conspecific with either amazonica and/or reticulata (both sensu this paper). However, based on our (J.L. Brown, E. Twomey, unpub. data) observations on Puente Itaya frogs (the type localiy of ignea ), we are confident the frogs described by Melin are referable to reticulata and not amazonica . The holotype (NHMG 512) possesses dorsolateral and middorsal stripes (partially broken) that extend about three-fourths of the SVL (14.9 mm), a characteristic common in striped R. reticulata . In contrast, in R. amazonica the middorsal stripe typically terminates between the shoulders (i.e., about one-fourth of the SVL). Lastly, we observed several populations of R. reticulata along the Iquitos–Nauta road, especially near Puente Itaya, that possess black dorsal markings similar to the holotype of ignea (which forms a dashed ‘U’). As a conclusion, we place Dendrobates tinctorius igneus Melin, 1941 as a junior synonym of Ranitomeya reticulata ( Boulenger, 1884 “1883”).

Distribution. This species in known to occur within the Amazonian rainforests of Peru (Loreto Department) and Ecuador (Pastaza Province), Fig. 21 .

Conservation status. Following the IUCN Red List categories and criteria ( IUCN 2010), we tentatively suggest listing this species as Near Threatened (NT). Although the distribution of this species is estimated to be around 20,000 km 2, much deforestation has occurred along Río Amazonas. Further, Iquitos, a very large city, occupies the center of its distribution.

Figures

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 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 21. Known distribution of Ranitomeya reticulata and R. ventrimaculata. 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).