Roosmalen et al., 2002
Type locality: West bank of the lower Rio Aripuanã, at the edge of the settlement of Nova Olinda, 41 km southwest of the town of Novo Aripuanã, Amazonas state, Brazil. This region is located in south-central Amazonia, south of Rio Amazonas and east of Rio Madeira. Coordinates for the type locality are: 05°30’63”S, 60°24’61”W. Altitude 45 m. The holotype is a complete adult skeleton and skull of unknown sex, found on the forest floor. The specimen apparently died from natural causes and was collected by M. G. M. van Roosmalen in November 1998. It was deposited as INPA no. 3929 in the Mammal Collection of the National Institute for Amazon Research (INPA), Manaus, Amazonas, Brazil. Paratypes: in 1996, two juvenile males were obtained alive along the Rio Mariepauá at Santa Cruz, not far from its confluence with the Rio Madeira, and were kept in M. G. M. van Roosmalen’s Breeding Center for Endangered Amazonian Monkeys in Manaus, Brazil. They died in April and August 2001 and are deposited as INPA no. 4029 and INPA no. 4033, respectively, in the Mammal Collection of the National Institute for Amazon Research (INPA), Manaus, Amazonas, Brazil.
Distribution: Interfluve delineated by the Rios Madeira-Jí-Paraná and Rios Aripuanã-Roosevelt, in the states of Amazonas and Rondônia, Brazil. In Rondônia, the species is parapatric in the west with C. brunneus along the entire Rio Jí-Paraná, and in the east with C. cinerascens along the Rio Roosevelt; in Amazonas, the species is parapatric with C. dubius in the west along the middle Rio Madeira, and with C. cinerascens in the east along the Rio Aripuanã. Ferrari et al. (1996) observed a grey titi monkey at Pimenta Bueno in Rondônia, on the west bank of the Rio Ji-Paraná. They noted that it was not the distinctively brown-coloured C. brunneus and were unable to identify it. It may have been C. bernhardi, which would extend its range a little to the west across the upper the Rio Ji-Paraná, but this requires confirmation. This species has been observed in the wild by M.G.M. van Roosmalen at the following localities: west bank of lower Rio Aripuanã, Nova Olinda, Amazonas state, 05°30’01”S, 60°24’27.4”W; west bank of lower Rio Aripuanã, Monte Alegre, Reserva Florestal Getal, 05°34’68”S, 60°23’40”W; west bank of lower Rio Aripuanã, Novo Oriente (Capimtuba), 05°43’41”S, 60°17’09”W; east bank of middle Rio Madeira, seringal São Luis, ca. 5 km south of the town of Manicoré, 05°50’28”S, 61°18’19”W, altitude 45 m.
Description: Upper and outer surface of head, trunk, and limbs greyish black, on the back mixed with brownish agouti or red-brown; forehead not defined from crown, greyish black to grey; ears black with conspicuous whitish tufts; sideburns, under parts of body, and inner side of limbs sharply contrasted dark orange; cheiridia sharply contrasted white against greyish black lower limbs; tail black except for a sharply contrasted white pencil.
Distinguished from C. cinerascens by uniformly dark orange instead of greyish sideburns and inner sides limbs, chest, and belly, and tail with white pencil; from C. brunneus by greyish instead of dark brown to black forehead and crown, and dark orange sideburns, inner sides of limbs, chest, and belly; from C. hoffmannsi and C. baptista by strikingly contrasting white ear tufts, cheiridia and tip of the tail (pencil); from C. moloch by greyish forehead and crown, white ear tufts, and blackish tail with a distinct white pencil; from C. dubius by lack of black vibrissae and white blaze.
Description of holotype: Forehead, crown, sides of body, and outer sides of limbs greyish; rump, mid-dorsum, back, and nape greyish mixed with brownish agouti to reddish brown, the hairs 5 cm long, with 5 blackish bands alternating with 4 narrow brownish agouti to red-brown ones, the most proximal (2 cm long) and the distal one (tip) black; face black with some white hairs around mouth and nostrils; ears black with white tufts contrasted with light greyish forehead and crown; tail ca. 55 cm long, the distal 7 cm forming a white pencil.
Measurements: See publication here.
van Roosmalen, G.M.; van Roosmalen, T. and Mittermeier, R.A. (2002). A taxonomic review of the titi monkeys, genus Callicebus Thomas 1903, with the description of two new species, Callicebus bernhardi and Callicebus stephennashi, from Brazilian Amazonia. Neotropical Primates 10(Suppl.): 1-52.
Locality: a 50 ha reserve adjoining the Floresta Amazonica Hotel in Alta Floresta, about five minutes from the airport, Mato Grosso, Brazil.
Almeida Noronha et al., 2007
Remarks: During this study we found no overlap between the range of C. cinerascens range and that of any other species of Callicebus, suggesting that this species is parapatric with its sister taxa, C. hoffmannsi, C. baptista and C. bernhardi.
Reis Monçâo et al., 2008
Distribution: West of the Río Yparaná, Alto Alegre dos Parecis (12º07’41’’S, 61º51’02’’O), Pimenta Bueno (11°36’30”S, 61°09’49”O) and Cacoal (11°24’13”S, 61°27’47”O).
Localities: see publication.
Description: Pelage chromogenetic analysis shows C. moloch has great color tone variation on several chromogenetic fields, especially on the ventral surface, which ranges from yellow to reddish-brown. I could split the specimens into three phenotypes: “normal phenotype”, “red phenotype” and “light phenotype”.
The “normal phenotype” is the commonest (84% of the sample) and has a cream forehead, crown (banded hair showing light bands broader than dark ones) flanks, dorsal surface of limbs, feet and hands; lower-back light brown with a slight brown stripe along the middle back, slightly darker than the flanks, not washed with brown or it has very little amount of this pigment. The middle portion of tail is very dark (from dark brown to black) and the tip lightening to very light brown or dirty white. Beard, chest, belly and ventral surface of limbs are light orange-brown, more pigmented at the tip of hairs.
The general color pattern of all specimens follows the description above, but specimens IPBHN 207, 208, 209 (loc. 52, Ig Almas, Rio Juruena, extreme north of Apiacás, MT); MZUSP 18956 (loc.53 – RO, Nova Colina Polonoroeste); MZUSP 18964, 20253, 20255, 20058, 20067 (loc.54 – RO, Nova Brasília Polonoroeste); MPEG 21972 (loc. 112 – PA, Ig. do Patauá, Município de Itaituba); MPEG 22000 (loc. 113 – PA, Apui, BR-230 Humaitá-Itaituba km 17) have the ventral pelage extremely pheomelanized of a live reddish-brown. These represent what I called “red phenotype”.
A third phenotype, called here “light phenotype” has ventral parts much lighter, sort of a lime-yellow (specimens MZUSP 5198 and 5200 from loc. 82 – AM, Bom Jardim, right margin of Amazonas River); MPEG 22014, 22015, 22016, 22017 (loc. 109 – PA, UHE Tucuruí, Tocantins River); MPEG 245 (loc. 95 – PA, São João do Araguaia); MPEG 246 (loc. 94 – PA, Alto Iriri River, Xingu).
Roosmalen et al. (2002) described C. bernhardi and identified specimens MPEG 22996, 22997 (locality 50 – BR km 150 Apis-Humaitá, right margin of Marmelos River, AM); MPEG 24590 and 24591 (locality 55 – Alta Floresta, MT) as belonging to this taxon. Paratypes of C. bernhardi (INPA 4029 and 4033; locality 57 – AM River Mariepauá left aff. River Madeira) show the same chromogenetic pattern as C. moloch, with identical chromogenetic fields. These specimens differ only in color tone and pigment amount on the ventral surface, exactly as seen in the “red phenotype”. In Roosmalen et al. (op. cit.), diagnostic characters that distinguish C. bernhardi of C. moloch are described as follows: “…by grayish forehead and crown, white ear tufts, and blackish tail with a distinct white pencil ”. Actually, there is wide variation in forehead and crown color tone among all 183 specimens of the 3 phenotypes, from grayish to light red-brown, and the description above agrees perfectly with most specimens analyzed of “normal phenotype” as well. Concerning the auricular tufts, none of 183 specimens of C. moloch (3 phenotypes) and those identified as C. bernhardi in INPA and MPEG that I could analyze, presented white auricular tufts (including C. bernhardi paratypes). Tails of all “red phenotype” specimens as well as C. bernhardi specimens are identical to C. moloch: black with a lighter tip. Drawings of C. moloch in Roosmalen et al. (2002) do not show a black tail and the whitish back of the hands, not matching all specimens analyzed. Thus, all specimens of the “normal phenotype”, “red phenotype”, “light phenotype” and those described as C. bernhardi show the same chromogenetic field pattern, differing, as mentioned, only in the amount of pigment (color tone) of the ventral surface.
Concerning the geographic distribution of C. moloch (all phenotypes), it is the broadest among all Callicebus species, occurring south of the Amazonas River, between the right margin of Madeira/Ji-Paraná Rivers to the left margin of Tocantins River. C. moloch is not found between the right margin of Aripuanã River and the left margin of Abacaxis River, where C. cinerascens is found (Noronha, et al. 2007). Callicebus moloch is found in Rondônia on both margins of the medium/upper Ji-Paraná River (Ferrari, et al. 2000), what is confirmed by specimens MZUSP 18956 (RO, Nova Colina Polonoroeste, right margin of Ji-Paraná River 10°48’S61°43’W, “red phenotype”; MZUSP 18964, 20253, 20255, 20058, 20067 (RO, Nova Brasília Polonoroeste, right margin of Ji-Paraná River – 10°56’S61°20’W “red phenotype”, and MPEG 19709, 19710, 19712, 19713 (Alvorada d’Oeste, BR 429 linha 64 km 87, left margin of Ji-Paraná River – 11°23’S62°18’W normal phenotype. Monção et al. (2008) also assigned specimens they called C. bernhardi (here, “red phenotype”) to 90 km west of Alto Alegre dos Parecis (Chapada dos Parecis, Rondonia). Roosmalen et al. (2002) states that there is a gap in the range of Callicebus at the southern portion of this region, between Sucunduri/Juruena River and Tapajós River. I could not find any specimens in Brazilian museums from this region. Wide rivers such as the Juruena / Teles Pires / Tapajós are no barriers isolating the three phenotypes of C. moloch. Gascon et al. (2000) observed that wide rivers are not always obstacles to put apart small mammals and frogs as well. Localities for C. bernhardi indicated by Roosmalen et al. (2002) are: 51 (AM, Comunidade de Nova Olinda, right margin of Aripuanã River, Novo Aripuanã – holotype, INPA 3929 only skeleton) and 57 (AM, Mariepauá River, right tributary of Madeira River – paratypes of C. bernhardi). Specimens MNRJ 2480 and 2481 (from AM, right margin of São João do Aripuanã River) presents “light phenotype” and this locality is only 30 km straight line from locality 51 and 60 km from locality 57, mentioned above, on the same bank of Aripuanã River. In the locality 109 (PA, UHE Tucuruí rio Tocantins) it is possible to find both “light and normal phenotype” as can be seen in specimens MPEG 21442, 21443, 22014, 22015, 22016, 22017, 22016 (normal phenotype) MPEG 22018 (light phenotype), one evidence of polymorphism. “Red phenotype” can be found far to the east from known localities of C. bernhardi. Specimens MPEG 21972 (locality 112- Ig. Patauá, Itaituba, PA), MPEG 22000 (BR 230 Itaituba, PA) and IPBHN 207, 208, 209 (locality 52- Ig. Almas, Juruena River, Apiacás, MT) are “red phenotype” (see Appendix I for coordinates). These localities are among others where phenotype can be normal phenotype or light phenotype, one more evidence of polymorphism.
One specimen from Alta Floresta (locality 55) MPEG 24590, label identificated as C. bernhardi, had its DNAmt sequenced and it is more similar to the sequence of IPBHN 207 (from Apiacás, MT), both “red phenotypes”.
A phylogenetic analysis for Callicebus carried by me (to be published elsewhere) shows strong evidence for the three phenotypes of C. moloch to be considered a polymorphism of the same taxon. Also, C. bernhardi appears as sister group of C. moloch. It is possible to recognize a trend to a clinal variation along a east-west transect through the range of the species, with specimens from western localities showing more pigmented ventral parts (phenotype red) and specimens with lighter ventral parts (phenotype light) to the east. “Normal phenotype” is found throughout the range. Moore (2009) found similar results in C. cupreus. C. hoffmannsi showed similar south-north differences in ventral amount of pigments as can be seen bellow. Based on this, I suggest here C. bernhardi, Roosmalen et al. (2002), to be considered as a junior synonym of C. moloch.