Literature discolor

Geoffroy Saint-Hilaire and Deville, 1848

Callithrix discolor

Locality: Peru

Description: Pelage speckled grey above and on the outer side of the arms and thighs; grey-whitish on the front of the head; a vivid red-brown on the underside and almost the entire limbs. Tail ashy-grey, with the tip of the hairs white.

Remarks: Related, but different on several characteristics, especially the colour of the forehead and a shape of the inferior mandible, to a species of the same region that we include, not without doubts, to C. cupreus.

Geoffroy Saint-Hilaire, I. and Deville, E. (1848). Note sur huit espèces nouvelles de singes   américains, faisant partie des collections de MM. De Castelnau et Emilie Deville. Comptes    rendus hebdomadaires des séances de l’académie des sciences de Paris 27 : 498.

I. Geoffroy, 1851discolor geoffroy 1851 9x13

Callithrix discolor

Distribution: banks of the Amazon and the Ucayali, Brazil and Peru.

Description: Upper parts grey, more or less red and speckled, a vivid red-chestnut brown on the under parts and on almost the complete limbs; tail grey with the extremity of the hairs white. Like C. moloch, C. cupreus and C. donacophilus this species is covered on the upper parts with long annulated hairs, and on the under parts with shorter hairs of a uniform colour. This colour, in the species that I have mentioned, is more or less red-rusty; in none of them is it as rich as in C. discolor. In him the throat, the cheeks, the chest, the belly, the inner sides of the thighs and arms and the totality of the arms and forearms are a beautiful red-chestnut. On the upper parts of the body, on the outside of the thighs, the shoulders and the arms, the hairs are brown-blackish at their base and then coloured with small rings of whitish and black; this gives a grey-red speckled tinge. The upper side of the head is also, in some of our individuals, speckled grey-red, with the forehead blackish; in others a grey with a weak red hue, with only a few black hairs on the forehead. The face is naked and blackish, except for the contour of the mouth, which is covered with white hairs. The tail is, at its base, speckled grey-red, like the upper parts of the body. It then becomes brown speckled with white, the extremities of the hairs being white, then whitish. The hairs of the terminal part, yellow or yellowish at their base, are blackish in the intermediate zone, then whitish. That is at least the colouration of most of the hairs of the extremity of the tail; a small number has black tips.
The nice series of individuals that possesses the museum is owed to the voyage of Castelnau and Deville. Among these individuals we can find a very young subject; he differs only from the adults in have the upper parts more reddish, of which the hue approaches very much that of the under parts, the hairs not being speckled. We have noticed, among the adults, that there is a large contrast between the upper and under parts, hence the name discolor, that M. Deville and I gave to the species.
The resulting characters that it is well distinguished from both C. moloch as C. donacophilus: the red of the under parts is in them much lighter and more ‘golden’ than chestnut.
Concerning C. cupreus of Spix, it appears, according to the plate provided by the author, at least by its description, that it approaches C. discolor by the colouration of its under parts; but, according to Spix, it has the tail grey-reddish, and that character distinguishes it from C. discolor, as well as from C. moloch and C. donacophilus. The comparison of the bones confirms the specific distinction between C. cupreus and C. discolor.
All individuals originate from the banks of the Amazon or its affluent the Ucayali. Those who origin from the Amazon , either the Brazilian or Peruvian part, resemble each other very much; those, on the contrary, from the Ucayali, in the Sarayacu mission, present a difference in the colouration of the head; in them we find the crown grey, and not grey-red, and only a few black hairs on the forehead. One differs from the others by having the fingers partly whitish or light reddish. Because of the latter character we hesitated if the latter belonged to the same species; but we established the presence of some whitish hairs on the border of the nail in some other individuals with red-chestnut fingers.

Geoffroy Saint-Hilaire, I. (1851). Description des mammifères nouveaux ou imparfaitement connus de la collection du Muséum d’Histoire naturelle 3 : Famille des Singes, Supplément. Archives du Muséum d’histoire naturelle 5 :551-555 + plate 28.

I. Geoffroy Saint-Hilaire, 1851

Callithrix discolor

Synonym: C. discolor (Geoffroy and Deville, 1848).

Remarks: The Paris Museum has several specimens:

– a series of type specimens, one conserved in alcohol. All from the banks of the Amazon, Peru and Brazil, from where they were brought by Castelnau and Deville in 1847. All show much resemblance, with the exception of the very young individuals, which have darker under parts;
– one male and a female came from the same collectors, from the Sarayacu mission. They differ from the other individuals of this pretty species by their speckled clear-grey forehead; one of them has a part of the fingers whitish.

Geoffroy Saint-Hilaire, I. (1851).  Catalogue méthodique de la collection des mammifères. Part 1 Mammifères, Introduction et catalogues des primates pp. 39-41.

Wagner, 1855

Callithrix cuprea

Synonyms: Callithrix cuprea (Spix, 1823; Wagner, 1840, 1848); Callithrix discolor (I. Geoffroy, 1848, 1851, 1851).

Remarks: In the description of I. Geoffroy of C. discolor is nothing that cannot be seen in C. cuprea. One could think that the nice figure of Geoffroy was done after one of our specimens. The mistake, to propose an old species as a new one, could have been avoided if he, not satisfied by the description of Spix, had studied my remarks on C. cuprea of 13 years ago.


Castelnau, 1855

Callithrix discolor (I. Geoffroy and Deville)

Synonym: Callithrix discolor (I. Geoffroy and Deville, 1848; I. Geoffroy, 1851).

Description: Pelage upperparts greyish-red and speckled, under parts and almost the complete limbs very vivid red-brown; tail grey with the extremities of the hairs white. Like the C. moloch that M. de Hoffmansegg has showed us in 1807, and like the two elegant species that we know since more recently, the C. cupreus of Spix and C. donacophilus of M. d’Orbigny, our C. discolor is covered on the upperparts with long annulated hairs and on the under parts with shorter hairs of a uniform colour. This colour, in the species that I cited, is always more or less reddish; in none of them it is richer than that of C. discolor. In this species, the throat, the cheeks, the chest, the belly, the inner side of the thighs and the arms, and the totality of the arms and underarms, are a pretty red-brown. On the upperparts of the body and the outer sides of the thighs, the shoulders and the arms, the hairs are brown-blackish at the base, but are further coloured with small rings of yellowish and black; this results in a general grey-red speckled hue. The top of the head is also, in some of our individuals, a speckled grey-red, with the anterior part of the front blackish; with some others, a grey vaguely tinted with red, with only a few black hairs on the front. The face is naked and blackish, except for the mouth, which is covered with white hairs. The tail is, at the base, a speckled grey-red, as the upper parts of the body; it becomes then brownish speckled with white, the extremities of the hairs being white. The hairs of the terminal part, yellow or yellowish at their base, are blackish in the intermediate zone and then whitish. This is how the colouration of most of the hairs at the extremity of the tail is; only a small number of the hairs have a black point.
The size of the monkeys is 3 decimetres for the adults, excluding the tail, which is approximately 3.5 decimetres.
The nice series of individuals that is in the possession of the Museum origins from the voyage of MM. de Castelnau and Deville. Amidst the individuals is a very young subject; he doesn’t differ from the adult but for a colouration more reddish on the upper parts, of which the tinge, which the hairs that are not speckled, approaches the colour of the under parts. In the adults we have seen a very marked contrast; this is where the name discolor comes from, that M. Deville and I gave to this species.
The characteristics that result, for the C. discolor, from the preceding description distinguishes it clearly from C. moloch and C. donacophilus; the red of the under parts is in these species much lighter and more golden than brown. However, C. cupreus of Spix, according to the plate given by the author and his description, seems to approach C.discolor by its colouration of the under parts; but, after Spix, it has a grey-reddish tail, and this characteristic distinguishes it from C. discolor, C. moloch and C. donacophilus. The comparison of the skulls confirms the distinction between C. cupreus and C. discolor: the latter has the inferior side of the lower mandible more rounded at the back, and rectilinear. It is concave, according to the figure of Spix, in C. cupreus. In C. discolor, the zygoma is notable larger than in C. cupreus. However, M. Wagner considers that our C. discolor belongs to C. cupreus.

All discolors in possession of the Museum originate from the borders of the Amazon or its affluent the Ucayali. Those that originate from the Amazon, both the Brazilian and the Peruvian part of the river, resemble each other much. Those who, on the contrary, come from the Ucayali, from the mission of Sarayacu, present a difference in the colouration of the head; it is with them that we find the grey crown, and not grey-red, and only a few black hairs on the front. There is one that differs by having a part of the fingers whitish or clear reddish. Because of the latter characteristic, we have hesitated some time to consider this individual to belong to the same species; but we have established that some animals with red-brown fingers have a few whitish hairs around the nail. It is therefore only a difference in quantity.

Castelnau, F. de (1855). Animaux nouveaux or rares recueillis pendant l’expédition dans les parties centrales de l’Amérique du Sud, de Rio de Janeiro a Lima, et de Lima au Para; exécutée par ordre du gouvernement Français pendant les années 1843 a 1847. Anatomie pp. 10-13. 

Dahlbom, 1857

Callithrix discolor

Synonym: Callithrix discolor (I. Geoffroy).

Locality: Peru.

Description: Hands with red hair. Cheeks, chin, throat, chest, abdomen, limbs inside and ends are rust-red. All other body-hair is black and yellow-brown or greysh, the tip of the tail is white-grey. Over the eyes is a black-brown stripe.

Measurements: body 370mm; tail 440mm.

Dahlbom. A.G. (1856-1857). Zoologiska Studier, afhandlande Djurrikets naturliga familjer 1: 151- 153.

Reichenbach, 1862discolor 9x13

Callithrix discolor

Synonyms: C. discolor (I. Goeffroy and Deville).

Distribution: Forests around the Amazon, Brazil and Peru.

Description: Upper side grey, more or less brown-red and speckled, underside and almost the complete limbs vivid chestnut-brown. Tail yellow-grey, towards the tip whitish.
One of the prettiest species of the genus. A young specimen is redder on the back, the same colour as the underside but speckled. The colour is supposed to mean that old specimens have much variation in colour.

Measurements: head and body 300mm; tail 350mm.

Reichenbach, H.G.L. (1862). Die Vollständigiste Naturgeschichte der Affen.

Gray, 1866

Callithrix cuprea (Spix, 1823) = Callithrix discors (sic) (Geoffroy)

Description: Fur soft, with abundant, elongated, stiffer hairs. The hands and feet red.

Gray, J.E. (1866). Notice of some new species of Callithrix in the collection of the British Museum. Annals and magazine of Natural History (3) 17: 57-58.

 Gray, 1870

Callithrix cuprea

Distribution: Brazil, Ega (Bates).

Synonyms: Callithrix cuprea (Spix, 1823); Callithrix discolor (I. Geoffroy, 1851).

Description: Dark black-and-grey grizzled; cheeks, throat, hands, feet, legs, and underside of the body dark red bay; tail like back, but rather darker; hairs of tail dark grey, with broad sub terminal black band; eyebrows black.

Variation: tail white at the end (Geoffroy).

Remarks: The figure given by Spix is very badly coloured, but the description agrees well with the specimens. Geoffroy’s figure is from a specimen with the end of the tail white; his figure is as much brighter than the Museum specimens as Spix’s is too sombre.

Gray, J.E. (1870). Catalogue of Monkeys, Lemurs and Fruit-eating Bats in the collection of the British Museum pp. 54-57.

Bartlett, 1871

Callithrix cuprea

Notes from Sclater:
I consider Wagner (Säugeth. V. p. 114) quite right in referring C. discolor of Is. Geoffroy to C. cuprea of Spix. Spix gives the Upper Amazon, frontiers of Peru, as its locality. Deville’s specimens of Callithrix discolor were obtained from Sarayacu and other places in the same district as that which Mr. Bartlett has collected in.

Bartlett, E. (1871). Notes on the monkeys from Eastern Peru. Proceedings of the Zoological Society of London pp. 219-220.

Martinez Saez, 1873

Callithrix discolor (Geoffroy)

Description: The description of C. discolor given by Geoffroy and the corresponding figure doesn’t match completely the specimen that I have in sight (= in the Madrid Museum). The white spot on the forehead, that appears to be much more marked in being brownish-reddish for the rest of the occiput, the numerous white hairs on the hands and feet, the brownish-reddish spots of the anterior half of the forearms and legs, and the more white tinge of the tail are the principal differences that I observe in this specimen. Geoffroy noted that some individuals have grey hairs on the vertex and whitish hairs on the fingers.

Martinez Saez, F.  (1873).  Nota sobre mamíferos Americanos. Anales de Sociedad Espanola de Historia Natural 2: 243.

Schlegel, 1876

Callithrix cuprea

Synonym: Callithrix cuprea (Spix, 1823) C. discolor (I. Geoffroy and Deville, 1848; I. Geoffroy, 1851).

Remarks: ….. the figure and description of C. discolor attributes to C. cuprea de Spix, it is clear that the form can be rejected, following the example of Gray, even in the case that Is. Geoffroy has confused the two species, which is very well possible, knowing that the two individuals, obtained by Castelnau near Sarayacu on the margins of the Ucayali differ from the others by their speckled light grey front and the partially whitish hairs on the fingers of one…..

The Leiden museum has one specimen of the voyage of Spix.

Schlegel, H. (1876). Les singes, Simia. Muséum d’Histoire Naturelle de Pays-Bas 12 :230-241.

Meerwarth, 1897

Callithrix cuprea

Synonym: Callithrix discolor (I. Geoffroy, 1844)..

Distribution: Amazonia, Peru; Tefé, Equador.

Description: Under and upper side different colours. The under side is vivid coppery or pallid red. Tail with long hairs at the base.
Throat and ventral side, sides of the head, inner sides of the limbs, and in some cases also the outer sides an intense coppery.
The same colour extends also on the exterior of the limbs, with exception of the thighs. The hairs on the rest of the body black or yellowish or yellow-grey-reddish.

Meerwarth, H. (1897-1898). Simios (macacos) do novo mundo. Boletin do Museo Paraense de Historia Natural y Etn. 2: 121-154.

Trouessart, 1898-1899

Callithrix cuprea

Synonyms: cupreus (Spix, 1823; Schlegel, 1876; Thomas, 1880); discolor (I. Geoffroy, 1848, 1851, 1851; Castelnau, 1855).

Distribution: Amazonia of Peru, Rio Solimoes, Rio Ucayali; Ega, Rio Teffe, Rio Huallaga, Rio Copotaza, Ecuador.

Trouessart, E.L. (1898-1899). Catalogus mammalium tam viventum quam fossilium 1: 44-46.

Cabrera, 1900

Callithrix cuprea leucometopa subsp. nov.

Remarks: The skull of this monkey cannot be confused with that of C. ornata. I have studied only the skull of a female, which compared to that of the C. cuprea of Spix does not show other differences than those that will be the result of the young age of the specimen. The symphysis of the jaw is, viewed from the side, rounder, but this character has also been mentioned by I. Geoffroy in his C. discolor, which is nowadays considered to be a real C. cuprea.

Measurements: Body 370mm/340mm; tail 365mm/335mm.

Cabrera, A. (1900). Estudios sobre una colleción de monos americanos. Anales de Sociedad Espanola de Historia Natural 9 (29): 83-85.

Thomas, 1908

Remarks: The museum specimens of C. cupreus, coming from the Upper Pastasa River, the Ucayali, and the Jurua, all agree in having the crown of the head distinctively more fulvous or reddish than the back, and this agrees with the description by Spix of his cupreus (type-locality Peruvian Amazons), while the Ucayali specimens (collected by Bartlett near Sarayacu) may be taken as topotypes of Geoffroy’s discolor, usually and rightly considered as a synonym of Spix’s name.

Thomas, O. (1908). Four new Amazonian monkeys. Annals and Magazine of Natural History (8) 2: 88-90.

Elliot, 1913

Callicebus cupreus

Synonym: Callithrix cuprea (Spix, 1823; Wagner, 1840; Reichenbach, 1862; Gray, 1870; Bartlett, 1871; Schlegel, 1876; Thomas, 1880; Forbes, 1896); Callithrix discolor (I. Geoffroy, 1845, 1848, 1851; Castelnau, 1855; Wagner, 1855; Dahlbom, 1856; Reichenbach, 1862; Sclatter, 1871).

Remarks: The specimen in Paris Museum marked Callithrix discolor I. Geoffroy et Deville, type, cannot be separated from the present species. It is somewhat faded in the lighter colours, but otherwise resembles C. cupreus. There are several examples in the Munich Museum obtained by Spix and all marked “type”. From one of these my description was taken.

Elliot, D.G. (1913). A review of the primates 1: 234-257.

Rode, 1938

Callithrix discolor

The museum in Paris has 3 type-specimens:

Specimen 95a (667/517).  Holotype. Locality: South-America (Banks of Amazon); Sarajacu Mission.

(Callithrix cuprea, Spix 1823)

Remarks: Brought along with a series of other individuals by Castelnau and Deville in 1847. Adult, moderate condition, no skull.

Specimen 95b (668/518). Allotype. Locality: South-America (Rio Janeiro) (Callicebus.nl: the board indicates Rio Javari!).

Remarks: Brought along by Castelnau and Deville on 01 January 1847. Young female with offspring, reasonable condition. Skull in mounted specimen.

Specimen 95c (669/519). Paratype. Locality: South-America; Sarajacu

Remarks: Brought along by Castelnau and Deville. Adult, moderate condition, skull in mounted specimen.

Rode, P. (1938). Catalogue des Types de Mammifères du muséum national d’histoire naturelle – 1, Ordre des Primates pp. 34-36.

Lönnberg, 1939

Callicebus cupreus

Synonym: C. discolor (I. Geoffroy)

Remarks: The specimens from the left side of Rio Jurua and Rio Eiru display a rather great variation, but the majority is rather dark and resemble in this respect the figure communicated by Spix. They may thus be regarded as representing the typical form. They resemble almost still more (although sometimes darker) the figure and description communicated by Is. Geoffroy of his C. discolor, which generally has been considered as identical with C. cupreus of Spix. Of special interest is that C. discolor is said to vary with regard to the colour of the head so that some specimens have “the anterior part of the front blackish” whole some others have “only a few black hairs on the front”.
Thomas described in 1914 a titi monkey of this group, which he named C. toppini. This one differs, however, from our specimens by having the tail hairs “on the proximal two-thirds tipped with black” instead of “white or buffy as in other species of this group”. Identification with this one is thus excluded, but it has been mentioned, because it is said to have “a distinct frontal black band”, and when further discussing this, the author quoted declares that C. cupreus has “no black hairs on the forehead”. If this is correct (it may be observed that the coloured plate by Spix of his type of cupreus shows a well conspicuous black frontal band, although the black does not extend further upwards”), it is evidently a mistake to synonymies the names cupreus and discolor, although that has been done by the majority of later authors. If this should be erroneous, and discolor represents a separate species our specimens from the region at Rio Jurua belong, of course, rather to that. It is, however, most probable, that cupreus compromises of a number, more or less distinct local races or subspecies, which cannot be satisfactory separated before a greater material has been compared from different localities.

Lönnberg, E. (1939). Notes on some members of the genus Callicebus. Arkiv för Zoologi 31 (13):1-18.

Cruz Lima, 1945

Callicebus cupreus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Callithrix discolor (I. Geoffroy et Deville, 1848).

The identity of the species cupreus of Spix and discolor of I. Geoffroy and Deville is no longer questioned. The cranial differences pointed out by I. Geoffroy in the commentaries on the zoological results of the trip of Castelnau and Deville do not seem apt, especially since the comparison was based merely on the drawings of the skull given by Spix.

Cruz-Lima, E. da (1945). Mammals of Amazonia Vol. 1. General introduction and primates pp. 175-198.

Vieira, 1955

Callicebus cupreus cupreus

Synonyms: Callithrix cupreus (Spix, 1823); Callithrix discolor (I. Geoffroy, 1848).

Distribution: East of Peru; Brazil: Amazonas (Joao Pessoa; Santo Antonio, Rio Jurua; Rio Eiru).

Vieira, C. da C. (1955). Lista remissiva dos mamiferos do Brasil. Arquivos de Zoologia 8 (10): 375-379.

Cabrera, 1958

Callicebus cupreus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Cebus cupreus (Fischer, 1829); Saguinus moloch variation (Lesson, 1840); Callithrix discolor (I. Geoffroy, 1848); Callicebus ustofuscus (Elliot, 1907); Callicebus cupreus (Thomas, 1908); Callicebus paenulatus (Elliot, 1909); Callicebus cupreus cupreus (Cruz Lima, 1945); Callicebus cupreus ustofuscus (Cruz Lima, 1945); Callicebus cupreus paenulatus (Cruz Lima, 1945).

Cabrera, A. (1958). Catálogo de los mamíferos de América del Sur. Instituto Nacional de Investigacion de la Ciencias Naturales, Ciencia Zoologica, 4 (1): 137-142.

Hill, 1960

Callicebus cupreus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); C. discolor (I. Geoffroy and Deville, 1848).

Remarks: Individual variation, not associated with age or sex, very considerable. According to Lönnberg, most specimens (from Rio Jurua and Rio Eiru) closely resemble Spix’s figure, but still more closely they approach the figure and description of discolor, commonly regarded as a synonym of cupreus. The variability of head colour agrees with that given for discolor.

The type of discolor is an adult, and is definitely an example of cupreus.


Hershkovitz, 1963

Callicebus moloch discolor

Synonym: Callithrix discolor (I. Geoffroy and Deville, 1848); Callithrix cuprea leucometopa (Cabrera, 1900); Callicebus leucometopa (Cabrera, 1917); Callicebus cupreus leucometopus (Cruz Lima, 1945); Callicebus subrufus (Elliot, 1907); Callicebus paenulatus (Elliot, 1909); Callicebus cupreus napoleon (Lönnberg, 1922); Callicebus rutteri (Thomas, 1923); Callicebus oenanthe (Thomas, 1924); Callicebus moloch oenanthe (Cabrera, 1958); Callicebus gigot oenanthe (Hill, 1960); ?Saguinus moloch (Poeppig, 1839); Callithrix cuprea (Bartlett, 1871); Callicebus cupreus (Thomas, – not Spix- 1928); Callicebus cupreus cupreus (Sanborn, 1949).

Type locality: Peru, subsequently restricted to Sarayacu, Rio Ucayali, Loreto by I. Geoffroy (1851). Type in Paris Museum.

Remarks: The first published description of Callithrix discolor is of a single specimen explicitly from Peru and characterized by a pale grey forehead and bright chestnut-red crown. Subsequent descriptions of discolor by I. Geoffroy are of an unspecified number of specimens collected by Castelnau and Deville in Brazil as well as Peru. Those from the Amazon River in Brazil and Peru with reddish crowns (I. Geoffroy, 1855) are said to be very much alike and here referred to cupreus. Those from Sarayacu, Rio Ucayali, distinguished from the others by their grey crowns, and , in one specimen, by its whitish or tawny toes, agree with the original description of discolor. Sarayacu, therefore, must be taken as type locality and the name discolor is the earliest available for the grey-fronted titis of the Rio Ucayali, Rio Huallaga, north bank of the Maranon and eastern Ecuador.


Freese, 1982

Callicebus moloch

Remarks: No Callicebus moloch territorial vocalizations were heard in the study sites of the reserve. Guards of the reserve claimed that C. moloch is found only along the headwaters of the Samiria, considerably upstream from the highest point on the river reached during censusing. Consequently, we do not consider C. moloch to be within the region of the study area.

Of possible ecological significance is a gap in the geographical range of Callicebus moloch in the basin of the Samiria and Pacaya Rivers in the region bounded by the Huallaga and Marafion Rivers on the northwest and the Ucayali River on the southeast. Neither our work along the Samiria nor the extensive work by Neville et al. (1976) along both the Samiria and the Pacaya revealed any good evidence placing C. moloch in the region, despite the occurrence of several collection localities around the periphery of most the Samiria-Pacaya region (Hershkovitz, 1963).

Freese, C.H.; Heltne, P.G.; Castro R., N. and Whitesides, G. (1982). Patterns and Determinants of Monkey Densities in Peru and Bolivia, with Notes on Distributions. International Journal of Primatology 3(1): 53-90.

Hershkovitz, 1987

Callicebus cupreus discolor

Hershkovitz, P. (1987): The titi. Field Museum of natural History Bulletin 58(6): 11-15.

Hershkovitz, 1988

Callicebus cupreus discolor

Member of the moloch group.

Remarks: Modern east bank (of Ucayali) C. c. cupreus and west bank C. c. discolor are part of a single cline. Some adults (ca. 1:100) of either subspecies are indistinguishable from extremes or near extremes of the other. Juvenils of C. c. discolor which usually lack the frontal blaze are inseparable from C. c. cupreus. The intergrading individuals of either subspecies are scattered throughout their respective range. A definable geographic zone of intergradation does not exist where subspecies ranges are separated by a shifting boundary river.

Prototypical C. cupreus discolor must have ranged along anarrow forested zone from southeastern Peru through Amazonian Ecuador into southern Colombia at least to the Sierra de La Macarena in the upper Río Orinoco watershed.

Modern C. c. ornatus, separated from C. c. discolor by a swath no less than 350 km wide, still retains all specific markings including reddish sideburns, under parts, and trend towards pheomelanization of limbs. Affinity with nearest relative discolor is  shown by the whitish transverse frontal blaze, marbled tail, and other shared derived features. The crown, however, still dominantly agouti in discolor, is pheomelanized in ornatus. In contrast, outer surface of lower arms and legs, completely pheomelanized in discolor, retain in ornatus most of the common ancestral agouti.

Differentiation of ornatus from discolor has been divergent with respect to some characters, and allometrically clinal in others. Notwithstanding marked changes, morphological separation is not quite complete, and hardly or not in all juveniles, perhaps of recency of genetic isolation. This suggests that reproductive and social isolation between ornatus and discolor is also incomplete. The two forms interbreed in captivity and produce fertile offspring. Their karyotypes are the same. These considerations suggest that ornatus may be best treated as a subspecies within the species Callicebus cupreus.

Hershkovitz, P. (1988) Origin, Speciation, and Distribution of South American Titi Monkeys, Genus Callicebus (Family Cebidae, Platyrrhini), by Philip Hershkovitz . Proceedings of the Academy of Natural Sciences of Philadelphia 140 (1): 240-272.

Hershkovitz, 1990

Callicebus cupreus discolor

Member of the moloch Group.

Synonyms: Callithrix discolor (I. Geoffroy, 1848); Callithrix cuprea leucometopa (Cabrera, 1900); Callicebus subrufus (Elliot, 1907); Callicebus paenulatus (Elliot, 1909); Callicebus napoleon (Lönnberg, 1922); Callicebus rutteri (Thomas, 1923).

Type locality: Sarayacu, Rio Ucayali, Ucayali, Peru. Holotype originally in the Museum National d’Histoire Naturelle, Paris. The holotype is no longer in the museum collection (the type is still in the museum – Callicebus.nl).

Distribution: Upper Amazonian region, in Peru between the Rios Ucayali and Huallaga south of the Rio Maranon, and between the Rios Napo and Santiago north of the Maranon; in Ecuador from east of the Andean foothills to the Rio Napo-Aguarico basin, north to the Rio Putumayo and across into Colombia to the south bank of the Rio Guamues. According to Soini (1982, p. 40), Callicebus c. discolor is absent from the lower Rio Ucayali basin and the Samiria-Pacaya watershed. I observed no titis in the region during fieldwork in 1980. Hernandez-Camacho and Cooper (1976, p. 49) speculated on the possible occurrence of C. c. discolor between the east bank of the Rio Napo-Aguarico and the right bank of the Rio Putumayo south of the Rio Guamues.
True provenance of the specimens of Callicebus c. discolor in the American Museum of Natural History “collected” by the Olallas at “Boca Rio Inuya, Rio Urubamba” and “Lagarto, Alto Ucayali” requires confirmation. The Rio Inuya is on the east bank or wrong side of the Rio Ucayali boundary of the discolor range, as here understood. The specimens from Lagarto, Rio Ucayali, Alto may actually have been taken at or near that place on the west bank of the Rio Ucayali. They agree with specimens from lower down the river at Yurimaguas and the Rio Pachitea region.

Description: Forehead, as a rule, with whitish or buffy tuft or transverse band or blaze sharply contrasted with agouti of anterior portion of crown; sideburns, side of neck, lower arms, legs, cheiridia, chest, and belly reddish, sharply contrasted with agouti crown, back, sides of body, and tail; tail mixed brownish and buffy agouti but often with pencilled tip to as much as terminal third dominantly buffy.

Measurements: See publication.

Comparisons: Distinguished from Callicebus cupreus ornatus and C. oenanthe by entirely reddish forearms and cheiridia; from C. caligatus and C. brunneus by whitish or buffy frontal tuft or transverse band; from C. caligatus, C. brunneus, and C. dubius by buffy, orange, or reddish agouti anterior portion of crown, upper surface of cheiridia reddish; from all other forms of Callicebus by one or more of above traits. The sole feature that almost consistently separates Callicebus cupreus discolor fromC. c. cupreus is the buffy or whitish tuft or transverse frontal blaze. In one of 5 specimens from Rio Tigrillo, Peru, the discolor blaze consists of a few dominantly whitish hairs; in another it is absent. In other samples from the same localities the blaze is well developed. In C. c. cupreus the blaze is normally absent, but in two individuals from Cerro Azul a small tuft of mid-frontal hairs are dominantly whitish. The blaze effect is created here by two narrow sub terminal pheomelanin bands of frontal hairs; in discolor, the effect is produced by a single wide sub terminal pheomelanin band of frontal hairs.

Specimens Examined: Total 133. Ecuador – Napo: Rio Coca, mouth; Rio Napo (2 syntypes of leucometopa); Rio Payamino; Rio Suno, mouth; San Francisco; Pastaza: Andoas (holotype of paenulatus); Montalvo; Rio Copataza; Rio Pindo; Not located: Rio Liguino. Peru – Amazonas: Bushimkin; Ciudad Constitution, Rio Palcazu; Kusu; Pagaat; Rio Maranon; Rio Santiago; Shimpunts; Tseasim; Huanuco: Tingo Maria; Loreto: Boca Rio Curaray; Iquitos; Lagartococha; Mishana, Rio Nanay; Mucha Vista, Rio Curaray; Puerto Indiana; Rio Tigrillo; Santa Luisa, Rio Nanay; Pasco: Puerto Leguia (holotype of rutteri); Puerto Victoria; Rio Pachitea (holotype of subrufus); Ucayali: Cumaria; Lagarto; “Rio Inuya,”; Yarinacocha.

Hershkovitz, P. (1990). Titis, New World Monkeys of the genus Callicebus: A Preliminary Taxonomic Review. Fieldiana Zoology 55: 1-109.

 Aquino and Encarnacion, 1994

Callicebus cupreus, two subspecies: cupreus and discolor

Description: The subspecies C. c. discolor has a wide transverse band of short, creamy-white hairs on the forehead; in C. c. cupreus the forehead is dark-brown. Total length up to 85cm and weight of 1200-1500g.


Kobayashi, 1995

Callicebus cupreus discolor

Based on cranial measurements, the genus can be divided in five groups:

  • the donacophilus group (including modestus, olallae, d. donacophilus and d. pallescens)
  • the cupreus group (including caligatus, c. cupreus, c. discolor and c. ornatus)
  • the moloch group (including brunneus, h. hoffmannsi, h. baptista, moloch and cinerascens)
  • the personatus group (including p. personatus, p. nigrifrons, p. melanochir)
  • the torquatus group (including t. lucifer, t. lugens, t. medemi, t. regulus, t. purinus and t. torquatus).

The group position of C. dubius remains uncertain; C. oenanthe and C. barbarabrownae were not examined.

A phylogenetic study of Titi Monkeys, Genus Callicebus, based on cranial measurements: 1. Phyletic groups of Callicebus. Primates 36(1): 101-120.

Hutterer et al., 1995

Callicebus cupreus discolor

Locality: Panguana, Peru

We follow Hershkovitz (1990) in recognizing cupreus as a species. A voucher specimen (SMNK) from Panguana fits the description of the subspecies discolor rather well. However, the ears of this animal are clearly white.

Hutterer, R., Verhaagh, M., Diller, J. and Podloucky, R. (1995). An inventory of mammals observed at Panguana Biological Station, Amazonian Peru. Ecotropica 1 (1): 3-20.

Brooks and Pando-Vasquez, 1997

Callicebus cupreus discolor

Locality: Sucusari Tributary, off the north bank of the Rio Napo (3°15’s, 73°05’W).

Remarks: ”our observations suggest that C. c. discolor is breeding north of the Rios Napo and Putumayo, on the Colombian border.

Brooks, D.M. and Pando-Vasquez, L. (1997). Crossing the great barrier: Callicebus cupreus discolor north of the Napo River. Neotropical Primates 5(1): 11.

 

Sheth et al., 2009

Callicebus discolor

Locality: Tiputini Biodiversity Station (TBS) in the Orellana Province of eastern Ecuador (∼0◦38’S, 76◦ 08’W)

Sheth, S.N.; Loiselle, B.A. and Blake, J.G. (2009). Phylogenetic constraints on fine-scale patterns of habitat use by eight primate species in eastern Ecuador. Journal of Tropical Ecology 25:571–582.

Groves, 2001

Callicebus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Callithrix caligata (Wagner, 1842); Callithrix discolor (I. Geoffroy and Deville, 1848); Callithrix castaneoventris (Gray, 1866); Callithrix cuprea leucometopus (Cabrera, 1900); Callicebus subrufus (1907); Callicebus ustofuscus (Elliot, 1907); Callicebus egeria (Thomas, 1908); Callicebus paenulatus (Elliot, 1909); Callicebus toppinii (Thomas, 1914); Callicebus cupreus napoleon (Lönnberg, 1922); Callicebus rutteri (Thomas, 1923); Callicebus cupreus acreanus (Vieira, 1952) and Callicebus dubius (Hershkovitz, 1988).

Description: Hairs have a long maroon-brown base, a straw-coloured band, a black band, another straw band, and sometimes a black tip. Hands and feet red. Tail brindled (hairs have straw-coloured base, long blackish shaft, straw tip). Crown agouti, becoming black anteriorly for various distances, but including forehead; sideburns reddish or orange; a white brown band variably present. Underside sharply marked reddish or orange, this tone extending to sides of the neck and inner surface of limbs; hands and feet reddish to whitish.

Remarks: Hershkovitz (1990) recognized four different taxa here: C. cupreus cupreus, C. cupreus discolor, C. caligatus and C. dubius. The first two were said to be predominantly phaeomelanic, the second two more eumelanic; in C. c. cupreus and C. caligatus there is no frontal band or just a small, agouti median tuft; in C. c. discolor and C. dubius there is a variable developed white frontal blaze.


Roosmalen et al., 2002

Callicebus discolor

Type locality: Sarayacu, left bank of Río Ucayali, Ucayali, Peru. The holotype is a skin and skull of the holotype, collected in 1847 by Comte Francis de Castelnau and Émile Deville, originally deposited in the Musée National d’Histoire Naturelle, Paris, France, but no longer in the museum collection. (= incorrect – callicebus.nl).

Distribution: Upper Amazonian region in Peru, south of the Río Marañon in the entire interfluve delineated by the Ríos Ucayali and Huallaga, and north of the Río Marañon between the Ríos Napo and Santiago; in Ecuador from the Andean foothills east to the Río Napo/Aguarico basin, and north to the Río Putumayo, and in Colombia to the right bank of the Río Guamués (Hernández-Camacho and Cooper,
1976). Brooks and Pando-Vasquez (1997) recorded this species just to the north of the Río Napo on the left bank of its northern tributary, the Rio Sucusari, in Peru. Hernández-Camacho and Cooper (1976) recorded its range in the Colombian trapezium, between the Ríos Putumayo and Amazonas. In Figure 1, C. discolor is given as occurring throughout the interfl uvium between the Ríos Putumayo-Içá and Amazonas-Solimões extending into Brazil, although this has yet to be confi rmed. In the eastern part of its range the species is parapatric with Callicebus cupreus along the Río Ucayali. Moynihan (1976, p.75) mentions the presence of titi monkeys of the moloch group in the close vicinity of the town of Valparaiso, between the Ríos Caquetá and Orteguaza in Colombia. These “lacked the white stripe above the eyes that is typical of both ornatus to the north and discolor to the south,” and Moynihan indicated that they could be an unnamed subspecies.

Description: Forehead with white or buffy tuft, contrasting with dark-brown transverse band, or blaze, this sharply contrasting with reddish crown and sideburns; often small white patches present alongside the lower jaw sharply contrasted with the reddish sideburns; sideburns, crown, side of neck, forearms, lower legs, cheiridia, chestand belly reddish, sharply contrasted with agouti back, sides of body, and tail; tail mixed brownish and buff-agouti, distal one-third to three-quarters predominantly buffy or white. Distinguished from Callicebus oenanthe and C. ornatus by entirely reddish forearms and cheiridia, from C. cupreus by one-third to three-quarters of tail buffy, white or buffy frontal tuft, and dark-brown blaze or transverse band.

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.

 Heymann et al., 2002

Callicebus cupreus

Locality: Along the Río Curaray, north-eastern Peru.

Heymann, E.W.; Encarnación C. F and Canaquin Y. J.E. (2002). Primates of the Río Curaray,   Northern Peruvian Amazon. International Journal ofPrimatology 23(1): 191-201.

 Rowe and Martinez, 2003

Callicebus discolor

Localities: south side of the Rio Aguarico, Cuyabeno NP, Ecuador (unconfirmed reports). Yasuni NP, south side of the Rio Aguarico (00°41.670’S, 076°27.736’W, 248m).

Rowe, N. And Martinez, W. (2003). Callicebus sightings in Bolivia, Peru and Ecuador. Neotropical Primates 11(1): 32-35.

Defler, 2004

Callicebus cupreus discolor

Distribution: C. c. discolor is found in Putumayo Department between the Guamués and Sucumbio rivers and may extend (acordign to some comments) to the Colombian trapezium south of the R. Putumayo (the latter needs to be confirmed and this author, despite efforts, has been unable to do so).

Description: A white band above the eyes. Above the white band another dark band (very contrasting); the wrists and the lower parts of the legs are reddish, as well as the feet and hands (not white).

Defler, T.R. (2004). Primates of Colombia.

Bueno et al., 2006

Callicebus cupreus cf. discolor

The male has a karyotype of  46 chromosomes (2n = 44; NF = 60: 16 Bi, 28 Acro).

Description: a male (from the Piscilago Zoo), with the chromatic characteristics agree with Callicebus cupreus cf. discolor, although the dark transversal band on the front is missing.

Bueno, M.L.; Ramírez-Orjuela, C.; Leibovici, M. and Torres, M. (2006) Información cariológica del género Callicebus en Colombia. Revista de la Academia Colombiano Ciencia 30 (114): 109-115. 

 Aquino et al., 2007

Callicebus discolor

Locality: watershed of the Río Alto Itaya (allong the quebradas Seis unidos, Agua Blanca, Nauta, Maquizapa, Miraflores and Yanayacu.

Aquino, R.; Pacheco, T. and  Vasquez, M. (2007). Evaluation and economic valorisation of the wild fauna in the Algodon River, Peruvian Amazonia. Revista Peruana de Biologia 14(2): 187- 192.

Vermeer et al., 2011

Callicebus discolor

Remarks: Callicebus discolor is often considered to be a species with red underparts and a large band of white hairs above the eyes (Aquino & Encarnacion 1994; Hershkovitz 1990; van Roosmalen et al. 2002). Isidore Geoffroy Saint-Hillaire and Deville described the species in 1848, based on monkeys Castelnau and Deville collected on the banks of the Amazon and Ucayali rivers (Geoffroy Saint-Hilaire & Deville 1848). Their first description of the new species caused considerable confusion, as they mentioned that it had some graywhitish hairs on the forehead. However, in a subsequent and more detailed description with a colored plate, Geoffroy reported the species to have a black forehead, in some cases “grey with a vague reddish hue” (Geoffroy Saint-Hilaire 1851). Personal examination by Jan Vermeer of the type specimens, present in the Paris Natural History Museum, showed that the species has no white band above the eyes, and is identical to the earlier described Callicebus cupreus (von Spix 1823). Other authors had already mentioned this synonymy (Elliot 1913; Gray 1870; Schlegel 1876), but it seems to have been ignored by most recent taxonomic revisers (Hershkovitz 1990; van Roosmalen et al. 2002).

Although the next available name for white-browed titi monkeys would be Callicebus leucometopus (Cabrera 1900), we need to determine first the validity of 5 other white-browed titi monkey species that have been described before we assign this name to the monkeys in the San Martin Department.


Aquino et al., 2014

Callicebus discolor

Localities: the survey area was on both sides of the road between Iquitos and Nauta, from appr. km 16 north till appr. km 85 south and from appr. 10 km to the east, limited by the Lower Itaya and the Amazonas and appr. 20 km to the west, limited by the Reserva Nacional Allpahuayo – Mishana (RNAM), quebrada Nauta and the upper parts of the quebradas Pensión and Miraflores.

Aquino, R.; López, L.; García, G.; Arévalo, I. and Charpentier, E. (2015). Situación actual de primates en bosques de alta perturbación del nororiente de la Amazonía peruana. Ciencia amazónica (Iquitos) 5(1): 50-60.

Aquino et al., 2015

Callicebus discolor

Locality: the forest in the lower basin of the Río Tigre and sub basin of the Río Nanay: Alto Itaya (UTM 615679/9540649), Alto Nanay (UTM 542669/9617188) and Huanganayacu (UTM 473383/9627228).

Rolando Aquino, R.; López, L.; García, G.; Arévalo, I. and Charpentier, E. (2015). Situación actual de primates en bosques de alta perturbación del nororiente de la Amazonía peruana. Ciencia amazónica 5(1): 50-60.