Ranitomeya imitator Schulte 1986

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: 68-72

publication ID

http://doi.org/ 10.11646/zootaxa.3083.1.1

persistent identifier

http://treatment.plazi.org/id/1D338788-9503-151C-C8FC-9BA63C13FC80

treatment provided by

Felipe

scientific name

Ranitomeya imitator Schulte 1986
status

 

Ranitomeya imitator Schulte 1986  

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

Figs. 3 View FIGURE 3 , 4 View FIGURE 4 , 9 View FIGURE 9 , 23 View FIGURE 23 (l–v), 24 (a–y), 25 (a–i), 28, 30

Tables 1, 4 – 6

Dendrobates quinquevittatus   (non Steindachner 1864)—Silverstone 1975 (partim): p. 35; Zimmermann and Zimmermann 1984 (partim): p. 35; 1988 (partim): p. 132

Dendrobates ventrimaculatus   (non Shreve 1935)— Caldwell & Myers 1990 (partim): p. 18, Fig. 11 View FIGURE 11

Dendrobates imitator Schulte 1986: p. 11   , Figs. 1–10 [MUSM BATR 10501 (holotype) collected by Rainer Schulte at km 33, near the village of San Jose, Carretera Tarapoto–Yurimaguas, San Martín, Peru];— Caldwell & Myers 1990: p. 17, Fig. 11c View FIGURE 11 ; Divossen 1999: p. 59; Symula et al. 2001: p. 2145, 2003: p. 452, Table 1, Figs. 1 – 6; Christmann 2004: p. 6, Figs. on p. 26, 27, 123 – 125; Brown et al. 2008a: p. 1140, Table 1, Fig. 1, 2008b: p. 1, Table 1, Figs. 1 – 3, 2009b: p. 478, Table 1 – 4, Fig. 3 View FIGURE 3 , 2009c: p. 148; Santos et al. 2009, by implication

Dendrobates imitator imitator   — Schulte 1999: p. 93 Figs. PB-015, PB-016; Lötters & Vences 2000: p. 253; Lötters et al. 2003: p. 1905; Christmann 2004: p. 142, Figs. on p. 142, 143

Dendrobates imitator intermedius Schulte 1999: p. 95   , Figs. ZB-009, PB-037, PB-034, PB-036, PB-029, PB- 040, PB-031 [CRS BD 27 (holotype) collected by Rainer Schulte at “Huallaga Canyon, Department San Martín, Perú, 200 m NN.”];— Lötters & Vences 2000: p. 252; Christmann 2004: p. 30, Figs. on p. 30, 127, 128

Dendrobates imitator yurimaguensis Schulte 1999: p. 104   , Figs. DB-056, PB-051, PB-035, PB-039 [CRS BD 41 (holotype) collected by Rainer Schulte on “Carretera Yurimaguas–Tarapoto, Alto Amazonas, Río Paranapura Drainage”];— Christmann 2004: p. 28, Figs. on p. 28, 29.

Ranitomeya imitator   — Bauer 1988: p.1; Grant et al. 2006: p. 171; Lötters et al. 2007: p. 478, Figs. 597–609; Brown et al. 2008c: p. 9; 2009a: p. 1877, Table 1, 2010: p. 436, Figs. 1, 2 View FIGURE 2 , 4 View FIGURE 4 , 5 View FIGURE 5 ; von May et al. 2008a: p. 396, Appendix 2

Ranitomeya intermedia   — Grant et al. 2006: p. 171; von May et al. 2008a: p. 396, Appendix 2

Dendrobates intermedius   — Santos et al. 2009, by implication

Background information. Few poison frogs have received as much attention from the public and the scientific community as R. imitator   . Initial studies were focused on this species’ validity and its mimetic relationship with R. variabilis   and R. summersi   , each sensu this paper, e.g., Schulte (1986), Symula et al. (2001, 2003). Recently, this species has been the focus of several studies regarding the evolution of parental care, mate choice and the evolution of Müllerian mimicry (e.g., Brown et al. 2008ab, 2009b, 2010).

This species was discovered in the late 1980s by Rainer Schulte (1986). After naming the species in 1986, Schulte (1999) further subdivided it into three subspecies, R. imitator imitator   , R. imitator yurimaguensis   and R. imitator intermedius   (described below as three color morphs). These names posed various taxonomic problems (see Lötters & Vences 2000) and immediately after their publication, the name imitator   yurimaguensis was formally synonymized with the nominotypical form by Lötters & Vences (2000). However, imitator   intermedius   tentatively remained. Grant et al. (2006) elevated the status of imitator   intermedius   to full species (as R. intermedia   ), but provided no justification for this taxonomic arrangement.

Ranitomeya imitator   has been the subject of several phylogenetic and population genetic studies and has been densely sampled throughout its known range ( Symula et al. 2001, 2003; E. Twomey & J.L. Brown, unpub. data). In addition, researchers have collected a considerable amount of acoustic, morphological and behavioral data on most populations of this species (E. Twomey, J.L. Brown & J. Yeager, unpub. data). None of the data from these studies indicate that R. imitator   is a species complex or adequately justifies the use of subspecies. In particular, recent population genetic studies have demonstrated that most color morphs are not reciprocally monophyletic, or even genetically distinct. Furthermore, within many populations different color morphs are known to coexist. To maintain the use of subspecies (and the resulting classification of R. intermedia   ) is misleading, ignoring considerable variation within and between most populations. Because of these reasons, we consider R. intermedia   to be a junior synonym of R. imitator   .

Definition and diagnosis. Assigned to the genus Ranitomeya   due to the combination of the following characters: adult SVL <20.0 mm, dorsal coloration conspicuous, dorsolateral stripes, when present, extend to top of thighs, brightly colored throat, distinctive pale reticulation on limbs and venter (absent in some populations), dorsal skin smooth, finger I greatly reduced and shorter than finger II, finger discs II – IV greatly expanded, disc of finger two times wider than finger width, thenar tubercle conspicuous, toe discs III – V moderately expanded, toe webbing absent, larval vent tube dextral, adults use phytotelmata for reproduction and deposit eggs away from phytotelm, maxillary and premaxillary teeth absent. Vocal slits present in males.

Rantiomeya imitator   is one of the most polymorphic poison frog species and a Müllerian mimic throughout most of its range, making identification of this taxon difficult. Furthermore, it possesses no single diagnostic morphological character that is consistent for all morphs. Three primary morphs exist: (i) Striped morph ( Fig. 24b–g, i– v View FIGURE 24 ), (ii) Spotted morph ( Fig. 23l–v View FIGURE 23 , 24a, h View FIGURE 24 ) and (iii) Banded morph ( Figs. 24w–y View FIGURE 24 , 25a–i View FIGURE 25 ). Although these three morphs predominate many populations do not exactly fit any of them (e.g., Fig. 24b, c, x, y View FIGURE 24 ). To thoroughly describe the morphs of a species that possesses such an immense variation undervalues complicated intrapopulation and interpopulation variation. Thus, the color morphs described below should be interpreted ‘loosely’ and in the context that they do not always represent a population or common ancestry.

(i) The Striped morph has a black dorsum with three thin yellow longitudinal stripes extending the length of the body. The presence of two paired spots on the nostrils creates the appearance of a yellow ‘cross’ anterior to the eyes (occasionally spots fuse to make a black “U,” Fig. 24o View FIGURE 24 ). Flanks are black and usually have a single yellow ventrolateral stripe that runs from the axilla to the groin. Legs, forelimbs and venter are black with fine light green to blue reticulation. The striped morph occurs throughout the lowlands northeast of the Cordillera Escalera, extending eastward, across the Río Huallaga , into the Pampas del Sacramento. It occurs as far north as Varadero (Department Loreto) but not reaching Río Marañón, and as far south as the northern Cordillera Azul, San Martín, Peru   .

(ii) The Spotted morph has a dorsal ground color of orange, golden yellow, or green. The dorsum has small to medium black spots that occasionally fuse, resembling irregular stripes. This morph typically possesses paired black spots on the nostrils (occasionally these spots are fused and resemble a ‘U’). Legs, forelimbs and venter are the same as the Striped morph, but can also be finely to coarsely reticulate in the colors of teal to gold. This morph is montane and occurs throughout the Cordillera Escalera as far north as Balsapuerto, Loreto, Peru and as far south as Chazuta, San Martín, Peru.

(iii) The Banded morph is the most variable; however, the majority of individuals possess a black dorsum with symmetrical orange (occasionally yellow) bands (occasionally possessing a complete or partial vertebral stripe). This morph is found throughout the central Huallaga canyon, San Martín, Peru.

Ranitomeya imitator   is a Müllerian mimic of R. summersi   (banded morph) and R. variabilis   (spotted and striped morphs, see discussion). Ranitomeya imitator   can be distinguished from these species by its loud, trill-like call (compared to faint buzz-calls in the model species) which is audible from over 5 meters. It can be distinguished from R. variabilis   by the presence of black paired nostril nose spots (single spot in R. variabilis   ). Ranitomeya imitator   can usually be distinguished from R. summersi   by the presence of a black ovoid head spot (versus pentagonal in R. summersi   ) and the absence of paired black gular spots (present in R. summersi   ). This species, particularly the Striped morph, is also similar in morphology to some morphs of R. sirensis   , although R. imitator   lacks a yellow ventral patch and white axillary and inguinal spots typical of most forms of R. sirensis   .

Tadpole. The description is based on a stage 26 tadpole from Cainarachi Valley, San José, San Martín, Peru. Body ovoid in dorsal view, wider near vent. Total length 17.5; body length 6.7; tail length 10.8, 62% of total length. Body width 3.7; body depth 2.2, 60% of body width. Eye well developed; naris small; distance from naris to anterior edge of eye 1.6. 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 1.7; tail muscle width at base of tail 0.9; maximum tail height 1.9. Dorsal fin slightly higher than ventral fin.

Oral disc ventral, weakly emarginate; transverse width 1.4, 38% of body width. Single row of small papillae present laterally and ventrally; dorsal gap where papillae absent. LTRF 2(2)/3(1) with A-1 developed on upper labium, A-2 with wide medial gap (one third total width of A-2); P-1 on lower labium with narrow medial gap; P- 3, 55% width P-1; P-2 equal in width P-1.

Color in preservative. Head whitish, mouthparts visible from above. Whitish abdomen, mostly transparent, intestinal coils black. Tail musculature uniform tan, dorsal and ventral fins opaque white.

Natural history. This species is a Müllerian mimic of several congeneric species throughout its range (see discussion). Ranitomeya imitator   exhibits a seasonal monogamous mating system and biparental care. After one to two embryos hatch, the male carries the tadpoles on its back and deposits each individually in a phytotelm. Both parents then return weekly and the female deposits trophic eggs after being stimulated by the male. The tadpoles are continually fed until metamorphosis. For details and discussion of these behaviors and their evolution, and the natural history of this species see Brown et al. (2008a, b, 2009b, 2010).

Vocalizations. The call of R. imitator   is stereotypical for members of the vanzolinii   group. It is a loud trill, notes 0.44 – 1.07 sec in length, repeated at 7 – 11 notes per minute. Despite major morphological differences between populations, acoustic divergence between populations appears to be minimal (E. Twomey, unpub. data).

Distribution. This species occurs within Amazonian rainforests of Peru (Departments: Loreto and San Martín), Fig. 30 View FIGURE 30 .

Conservation status. Following the IUCN Red List categories and criteria ( IUCN 2010), we tentatively suggest listing this species as Least Concern (LC). Its distribution is estimated to be around 10,000 km 2. Because of the pet market demand and extreme morphological variation over a small geographic area, some color morphs maybe vulnerable.

Kingdom

Animalia

Phylum

Chordata

Class

Amphibia

Order

Anura

Family

Dendrobatidae

Genus

Ranitomeya

Loc

Ranitomeya imitator Schulte 1986

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
2011
Loc

Ranitomeya intermedia

von May, R. & Catenazzi, A. & Angulo, A. & Brown, J. L. & Carrillo, J. & Chavez, G. & Cordova, J. H. & Curo, A. & Delgado, A. & Enciso, M. A. & Gutierrez, R. & Lehr, E. & Martinez, J. L. & Medina-Muller, M. & Miranda, A. & Neira, D. R. & Ochoa, J. A. & Quiroz, A. J. & Rodriguez, D. A. & Rodriguez, L. O. & Salas, A. W. & Seimon, T. & Seimon, A. & Siu-Ting, K. & Suarez, J. & Torres, J. & Twomey, E. 2008: 396
Grant, T. & Frost, D. R. & Caldwell, J. P. & Gagliardo, R. & Haddad, C. F. B. & Kok, P. J. R. & Means, D. B. & Noonan, B. P. & Schargel, W. E. & Wheeler, W. 2006: 171
2006
Loc

Dendrobates imitator imitator

Christmann, S. P. 2004: 142
Lotters, S. & Reichle, S. & Jungfer, K. - H. 2003: 1905
Lotters, S. & Vences, M. 2000: 253
Schulte, R. 1999: 93
1999
Loc

Dendrobates imitator intermedius

Christmann, S. P. 2004: 30
Lotters, S. & Vences, M. 2000: 252
Schulte, R. 1999: 95
1999
Loc

Dendrobates imitator yurimaguensis

Christmann, S. P. 2004: 28
Schulte, R. 1999: 104
1999
Loc

Ranitomeya imitator

Brown, J. L. & Twomey, E. & Pepper, M. & Rodriguez, M. S. 2008: 9
von May, R. & Catenazzi, A. & Angulo, A. & Brown, J. L. & Carrillo, J. & Chavez, G. & Cordova, J. H. & Curo, A. & Delgado, A. & Enciso, M. A. & Gutierrez, R. & Lehr, E. & Martinez, J. L. & Medina-Muller, M. & Miranda, A. & Neira, D. R. & Ochoa, J. A. & Quiroz, A. J. & Rodriguez, D. A. & Rodriguez, L. O. & Salas, A. W. & Seimon, T. & Seimon, A. & Siu-Ting, K. & Suarez, J. & Torres, J. & Twomey, E. 2008: 396
Lotters, S. & Jungfer, K. - H. & Schmidt, W. & Henkel, F. W. 2007: 478
Grant, T. & Frost, D. R. & Caldwell, J. P. & Gagliardo, R. & Haddad, C. F. B. & Kok, P. J. R. & Means, D. B. & Noonan, B. P. & Schargel, W. E. & Wheeler, W. 2006: 171
Bauer, L. 1988: 1
1988
Loc

Dendrobates imitator

Brown, J. L. & Morales, V. & Summers, K. 2008: 1140
Christmann, S. P. 2004: 6
Symula, R. & Schulte, R. & Summers, K. 2001: 2145
Divossen, H. 1999: 59
Caldwell, J. P. & Myers, C. W. 1990: 17
Schulte, R. 1986: 11
1986