Literature cupreus

Spix, 1823spix cupreus 9x13

Callithrix cuprea sp. nov.

Distribution: forest of Solimoes, Brazil.

Description: The coppery Sagoin, known by the locals as the ‘Yapusa”, is grey-brownish on the upper parts, and coppery on the under parts and the legs. The head is covered with short red hairs; the middle of the face is almost naked. The face is contoured from the temples to the lower cheeks with coppery coloured hairs, directed forwards. The lips and chin are covered with small black rigid hairs. The ears are naked on the backside, covered with some red hairs on the other side. Generally all hairs are directed backwards; those of the back, upper arm and thighs relatively long, black and on the same time mixed with red. The tail which is grey-reddish, becomes towards the tip paler and white. The eyes are brown.

Spix, J.B. de (1823). Simiarum et vespertiliarum Brasilienses species novea; … pp. 23.

Fischer, 1829

Cebus cupreus

Synonym: Callithrix cuprea (Spix, 1823).

Fischer, J.P. (1829). Synopsis mammalium Pp. 53.

Cuvier, 1829

Callithrix cuprea

Synonym: Callithrix cuprea (Spix, 1823)

Cuvier, G. (1829). Le Règne Animal, nouvelle édition 1: 104.

Cuvier and Voigt, 1831

Callithrix cuprea (Spix, 1823) is still an uncertain species.

Cuvier, G. and Voigt, L.S. (1831). Das Thierreich geordnet nach seiner Organisation 1: 96-97.

Wagner, 1833

Callithrix cuprea (Spix, 1823)

Remarks: Under the by Spix mentioned species of this genus is the C. cuprea the only new species, which characteristics would have been more obvious when the beautiful red colour of the under parts and the extremities on the plate had not become a dirty brick-red, and when the white colouration of the terminal halve of the tail clearer had been indicated.

Wagner, J.A. (1833). Critische Revision des brasilian. Affenarten. Isis von Oken 10(2):994.

Wagner, 1840

Callithrix cuprea

Synonym: Callithrix cuprea (Spix, 1823).

Locality: Solimoes, near the Peruvian border.

Description: The long hairs of the back are ringed with black and pale-yellowish, resulting in a mixed colouration, which becomes on the backwards directed hairs of the head reddish. Cheeks and whole underside of the body, the inner sides of the limbs, outer sides of forearms and lower legs till the tips of the fingers and toes, vivid coppery-red. The tail is below at the base pale-red; on the upper side at the base ringed black and red-brownish, but becomes in most individuals dirty-white, which is the dominant colour at 2/3 of the tail.

Measurements: head and body 350mm; tail 450mm.

Wagner, J.A. (1840). Schreber, die Säugthiere in Abbildungen nach natur mit Beschreibungen. Supplementband, Erste Abtheilung: Die Affen und Flederthiere Pp. 228-234.

Lesson, 1840

Saguinus moloch

Synonym: Cebus cupreus (Spix, 1823).

Variation A:

Distribution: Brazil, the forests of the Solimoes, on the border with Peru.

Thick pelage; naked face, blackish; a small line of white hairs on the upper lip; a moustache of some black hairs; back brown-grey; head reddish; cheeks, throat, chest; abdomen and hands coppery; tail greyish-red, ending in black; the other parts varying in brown, black or whitish.

Lesson, R.P. (1840). Species des mammifères bimanes et quadrumanes pp. 161-167.


Schinz, 1844

Callithrix cupreus

Synonym: Callithrix cuprea (Spix, 1823).

Distribution: Solimoes, on the border of Peru.

Description: The back has long hairs, the hairs are annulated black and red-yellow; the cheeks, undersides, inner sides of limbs, forearms and lower leg coppery-red. The tail is on the underside on the base pale red, on the upper side black, but becomes dirty-white towards the tip.

Schinz, H.K. (1844). Systematische verzeichniss aller bis jetz bekannte Säugethiere oder    Synopsis Mammalium nach dem Cuvier’schen System 1: 80-83. 

Boitard, 1848

Saguinus moloch

Synonyms: Saguinus moloch (Lesson, 1827); Callithrix moloch (I. Geoffroy, 1844); Cebus moloch (Hoffmannsegg, 1807; Fischer, 1829); Cebus cupreus (female) (Spix, 1823); Simia moloch (Humboldt, 1812).

Distribution: Brasil and Pará.

Remarks: One variety, the Cebus cupreus of Spix, Fischer and Cuvier, can be found in Brasil, in the forests of the Solimoes. It has a more luxurious fur; the face blackish; the head reddish; the cheeks, the throat, the chest, the belly and the hands coppery; a small white moustache on the upper lip; the back brown-greyish; the tail grey-reddish with the extremity black; the other parts are varied brown, black or whitish.

Boitard, P. (1848). Dictionnaire Universel d’Histoire Naturelle 11:290-292.

Lesson, 1848

Cebus cupreus

Distribution: Solimoes, on the border of Peru and Brazil.

Description: Brown back, head reddish and under parts and hands a coppery hue. The tails ends with black.

Lesson, R.P. (1848). Etudes sur les Mammifères Primates. Revue Zoologique par la Societé Cuvierienne 11:232-233.

Wagner, 1848

Callithrix cuprea

Synonym: Callithrix cupreus (Spix, 1823; Wagner, 1840).

Distribution: the forests on the Solimoes near the Peruvian border.

Wagner, J.A. (1848). Callithrix. Springaffe. Beitrage zur kenntniss der Säugthiere Amerikas Abhandlungen des Königl. Akademie des Wissenschaften München 5: 446-457.

Wagner, 1855

Callithrix cuprea

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

Distribution: on the Solimoes, near the Peruvian border.

Description: the upper side the shoulders and the largest part of the outer sides of the thighs are speckled with drab and black; the complete under side of the body and the inner sides of the limbs -with the exception of the inner sides of the forearms and lower legs- the hands and the cheeks are covered with uniform rusty coppery red hairs, that end in many specimens on the outside of the limbs with black tips. The drab rings of the hairs of the back are sometimes more towards red, sometimes more towards grey. The head is mostly reddish, sometimes it has the colour of the back. In some individuals is the black colour if the front of the head dominant over the drab. The tail, that has at the base the colour of the body, is speckled whitish, in some specimens the white is that present that it almost replaces the black, resulting in a tail that has the latter half almost dirty white.

Measurements: body 375mm; tail 450mm.

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.

Wagner, J. A. (1855). Schreber, die Saugethiere in Abbildungen nach der Natur mit Beschreibungen. Supplementband, Fünfte Abtheilung : Die Affen. 

Dahlbom, 1857

Callithrix discolor

Synonym: Callithrix discolor (I. Geoffroy, 1851).

Distribution: Peru.

Description: Hands reddish; under sides of body and inner sides of limbs red.

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

Reichenbach, 1862cupreus 9x13

Callithrix cuprea

Synonyms: Callithrix cuprea (Spix, 1823; Wagner, 1848); Saguinus moloch Var. A. (Lesson, 1840).

Distribution: Solimoes in Brazil, near the border with Peru.

Description: Back greyish black-brown, head reddish, cheeks, throat, breast, belly and all four legs coppery, tail at base reddish-grey, further black-brown or black, covered with white. Hairs on back, sides, upper arms, hips and tail sooty-black, towards the tip ringed black and whitish, almost 40mm long; shorter on skull, reddish or yellow-reddish; on the cheeks, sides of the neck, breast, belly and base of tail as well as the legs coppery; longer at the tail, on a third black-brown-grey, whitish towards the tip.

Reichenbach, 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

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

Distribution: Brazil, Ega (Bates).

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

Distribution: This monkey is equally widely distributed, but not so numerous, as the last named species (= Saimiri ustus); in fact it may be regarded as rather rare. I obtained specimens of it at Cashiboya on the Ucayali, and Santa Cruz on the Huallaga.

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.

Schlegel, 1876

Callithrix cuprea

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

Distribution: Spix has discovered this species on the margins of the Solimoes in Peru. Castelnau has observed it on the margins of the Peruvian Amazon and its affluent called Ucayali. Bates has obtained individuals shot near the Ega, on the margins of the Teffé and Bartlett, in small numbers, near Cashiboga on the margins of the Ucayali and near Santa-Cruz on the borders of the Huallaga.

Description: Coppery-red on the sides of the head, the underside of the throat and the body, as well as the four extremities with their hands, always with the exception of the outside of the thighs. Hairs of all other parts ringed with black and grey-yellow-reddish.

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.

Jentink, 1892

Callithrix cuprea

Remarks: the Leiden museum has four individuals:

– an adult female, one of the types of the species, from the banks of the of the Solimoes, Peru. From the collections of Spix;
– two adult individuals, from the Rio Copataza, Ecuador. From the collections of Buckley, 1877/78. Obtained from Frank in 1880;
– a semi-adult individual, from the Rio Copataza. From the collections of Buckley, 1877/78. Obtained from Frank in 1880.

Jentink, F.A. (1892). Catalogue systématique des mammifères (singes, carnivores, ruminants, pachydermes, sirènes et cétacés). Muséum d’Histoire Naturelle des Pays Bas 11: 51-53.

Forbes, 1896

Callithrix cuprea

Synonyms: Callithrix cuprea (Spix, 1823; Gray, 1870; Schlegel, 1876; Thomas, 1880), Callithrix discolor (I. Geoffroy, 1848, 1851; Wagner, 1855).

Distribution: This species is found throughout the whole of the Peruvian Amazons, though not in very large numbers – indeed, it is said to be rare. It has been recorded from Cashiboya on the Ucayali, and Santa Cruz on the Huallaga. Mr. O. Thomas mentions his having examined twelve specimens from the Copataza River, and one from Andoas in Ecuador. Of these he says: “The Andoas specimen, which is a male, differs from the rest in having the fur on the back of a dirty orange-grey colour, without annulations, instead of being of a bright annulated black and white. One of the others, a female, shows a tendency to this condition of the hair, which is, therefore, probably a seasonal change, as the Andoan specimen was shot in September, while the others were obtained between December and February”.

Description: Fur soft and woolly, mixed with numerous long stiff hairs; face black; back grizzly blackish-grey in colour; tail the same but darker; the basal part and tips of the hairs grey, with an intermediate band below the tips, black; tip of the tail sometimes white; the cheeks, throat, hands, feet, legs, and the underside of the body, dark reddish bay; the ears coppery-red.

Forbes, H.O. (1896). A Handbook of the Primates pp. 159-165 + plate 14.

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.

Ihering, 1904

Callicebus cupreus

Distribution: Rio Jurua.

Description: Dorsal side dark greyish, ventral side brown.

Ihering, H.  von (1904). O Rio Juruá. Revista do Museo Paulista 6 :412.

Trouessart, 1904-1905

Callicebus cuprea

Distribution: Amazonia of Peru; Ecuador (in mountains).

a) – leucomystax (Cabrera, 1900).

Distribution: Ecuador.

Trouessart, E.L. (1904-1905). Catalogus mammalium tam viventum quam fossilium. Quinquennale  Suppl. Pp. 25-26.

Elliot, 1907

Callicebus usto-fuscus

Type locality: “Brazil”

Description: Allied to C. cupreus, but much darker in colour; teeth much larger; palate longer and narrower; brain-case wider; space between pterygoid processes and bullae and the width of basioccipital throughout its length greater. Practically the skull is larger in every way and more massive. Mandible longer and heavier and the depth of the ramus greater.
General hue above burnt-umber, the hairs being slaty-grey at base, then annulated with two bands of slate- and two clay-colour, and a dark tip. Face naked, black; top of head a mixed dark ochraceous rufous and black, the black predominating on the forehead; the rump is redder than the back and is a burnt-sienna on the outer side of the limbs; hands and feet claret-brown; sides of head, throat, inner side of limbs, and under parts maroon; basal third of tail black, the hairs being chestnut with broad black tips, rest of tail mixed black and yellowish grey or very pale clay-colour, the underside of tail being almost altogether clay-colour; ears black.

Measurements: Size about the same as C. cupreus.

Skull: occipital region has been cut away; intertemporal width 32mm.; zygomatic width 41; palatal length 21; width between last morals 12; breadth of brain-case 35; length of nasals 9; length of upper molar series 14; length of m¹ 5; length of mandible 42; extreme height of mandible 35; length of lower molar series 17.5.

Remarks: B.M. no. of type: 51.7.3.1

This species is nearest C. cupreus, but is altogether different in colour and darker in all its hues. The skulls are also not at all in accord, the differences mentioned being very conspicuous when they are compared. The unique example has no history beyond the statement that it came from Brazil.

Elliot, D.G. (1907). Descriptions of apparently new species and subspecies of mammals belonging to the families Lemuridae, Cebidae, Callitrichidae, and Cercopithecidae in the collection of the Natural History Museum. Annals and Magazine of Natural History 7 (20): 191-193.

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

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

Type locality: Banks of the Solimoes River, Brazil (Munich Museum).

Distribution: Regions of the Peruvian Amazons; Solimoes, Ucayali, Huallaga and Copataza rivers; and Andoas, Ecuador.

Description: Face black; top of head grey, becoming orange rufous on occiput, or buffy-yellow on forehead grading into ferruginous on occiput, these colours due solely to the tips of the hairs which are black on the basal portion; upper parts reddish brown and black, the former being the tips of the hairs, producing an annulated appearance; sides of head, limbs, hands, feet, and under parts coppery red; tail mixed greyish white and black, the basal portion like the back; hairs on ears coppery red (ex. Type Munich Museum).

Measurements: Total length, 900mm; tail, 290mm; foot, 85mm.

Skull: occipito-nasal length, 56; zygomatic width, 39; intertemporal width, 36; palatal length, 18; width of braincase, 34; median length of nasals, 6; length of upper molar series, 14; length of mandible, 38; length of lower molar series, 15.

Remarks: There is some variation among individuals of this species, and some have the upper parts uniform Vandyke brown, palest on the centre of the back, without the annulations so characteristic of the typical style; the tail also is mixed ochraceous buff and black with a buffy tip, the rest of the pelage however, being coppery red as in the others.
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.

Thomas, 1928a

Callicebus cupreus

Locality: Cerro Azul, Contamana, 2000’ (female 1417).

Remarks: This species was also obtained by Bartlett at Cashiboya, East of Sarayacu, and at Santa Cruz, Huallaga. A series collected by Ehrhardt in the typical region of the Solimoes shows that the colour of the crown is so variable – rufous or grey – that C. egeria should probably be united with C. cupreus. The cranial differences described in C. egeria do not prove to be as constant as was supposed. C. paenulatus (Elliott) is also probably the same as C. cupreus.

Thomas, O. (1928a). The Godman-Thomas Expedition to Peru. – VII. The mammals of the Rio Ucayali.  Annals and Magazine of  Natural History (10) 2: 253-254.

Lönnberg, 1939

Callicebus cupreus

Synonym: C. discolor (I. Geoffroy)

Locality: The Stockholm Museum has 8 specimens from left (north-western) side of Rio Jurua, Joao Pessoa; 1 specimen from Rio Jurua, left side, Igarape do Gordao; 11 specimens from southern (right) side of the Rio Jurua and western (left) side of Rio Euru, near their junction, Santo Antonio.

Description: All these specimens agree in having the sides of the head, the throat, the lower side of the body, the limbs from elbow and knee and including hands and feet saturated rufous red in most cases resembling “mahogany red”, in others “chestnut” or even “maroon”. The top of the head is more or less rufous by means of such tips or subapical rings to the hairs, but the shade is varying in different individuals. Sometimes almost the whole head looks coppery red, in other cases the shade is more orange rufous in consequence of such tips or subapical rings to the hairs. There is always a black front-line against the naked face, but in many, mostly adult, specimens there is a rather broad shading of black upwards into the more or less mahogany red vertex. This colouration is, however, somewhat independent of age, because even a rather young female has a mahogany red head shading into black in front. The hairs on the ears are more or less dark reddish in correspondence with the colour of the other parts of the head. The colour of the back is very different in different specimens, because the hairs are in some annulated with rufous, in others with comparatively pale, more or less buffish rings. The most proximal portion of the tail has usually the same colour as the back, the remaining part has long dirty whitish tips to the hairs which, at least in the distal half, usually are quite dominating, but anteriorly along a certain extension les completely cover the blackish middle parts of the hair so that the tail is speckled.

Collectors measurements: Males: total length 790/790/712mm; tail 475/440/400mm; hind foot 95/92/92mm. Females: total length 765/730/715mm; tail 440/420/410mm; hind foot 100/95/90mm.

Skull: measurements and description of 6 individuals in publication.

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”. By this C. ustofuscus of Elliot with “black predominating on the forehead” appears to be included in the cupreus series as well, and the description by the author quoted of his only specimen from unknown locality suits also in several respects some of our specimens. 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.

Cabrera and Ypes, 1940

Callicebus cupreus

Distribution: Solimoes and Maranon and their affluents.

Description: Another species closely related to C. leucometopa and C. ornatus, and especially to the former. Differs principally by lacking the white band or diadem on the forehead. The dark brownish-grey, more or less washed with red on the upper parts; has the under parts, including the sides of the neck and throat, pretty dark coppery red, drawing sometimes towards chestnut.

Measurements: as the other species.

Cabrera and Ypes (1940). Mamiferos Sud Americanos. Compana Argentina de Editores, Buenos Aires pp. 85-88.

Vieira, 1944b

Callicebus cupreus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Callicebus cupreus (Ihering, 1904; Lönnberg, 1939); Callicebus cupreus cupreus (Lima, 1944).

Localities: Santa Cruz, Rio Eirú; San Antonio; Joao Pessoa, Rio Juruá.

Description: In our specimens we notice that some have the back more intense brown-reddish. Most variation is in the colour of the tail, some present almost white-yellowish in almost it whole length.

Vieira, C.C. (1944b). Nova Contribuiçao ao Conhecimento dos Mamiferos do Rio Juruá. Papeis Avulsos de Zoologia, Sao Paulo 4: 251-252.

Cruz Lima, 1945cruz cupreus 9x13

Callicebus cupreus cupreus

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

Distribution: The geographical range seems to be very wide. It is reported by the authors on the Peruvian and Brazilian Amazon, at least to Tefé. On the rivers Ucayali (Sarayacu, Cashiboya), Huallaga (Santa Cruz), Copataza, Pastaza (Andoas), Jurua, (Joao Pessoa, Igarape do Gordao, Santo Antonio, Euru River); and Aquiry or Acre, on the Brazilian boundary with Bolivia, Cerro Azul (Contamana).

Description: Face bare, black, with rare long stiff black hairs along the sides of the nose and on the chin, white fuzz on the upper and lower lips. A narrow frontal band above the eyebrows made up of black hairs with a mixture of some white and tawny ones; top of the head speckled, strongly washed with tawny, the hairs as a rule being light greyish brown at the base, this followed by three zones, one yellowish white, the second black and the last, apical, bright tawny, this being the most visible and giving the predominant colour of the region. On the nape, shoulders, back, outer surface of the thighs, loins and upper surface of the base of the tail the hairs have the same arrangement of colours which is observed on the crown, the yellowish-white band however, being the most extensive. On the flanks and thighs the general tone is paler, for the two terminal bands are less vivid than on the back. The body is less tawny than the head because on the much longer hairs of the body the lighter bands are more noticeable. Sides of the head in front of the ears and around the face and chin, the throat, forelimbs, hands, chest, under parts, inner sides of the thighs, legs and feet copperish chestnut red, darker at the wrist and ankles, with a mixture of blond hairs on the hands and feet. Upper surface of the base of the tail reddish; on the rest of the tail almost all the hairs have a yellowish basal portion followed by a brownish-black band and a pale tawny apical zone. Toward the end of the tail the hairs lose their dark bands, and the tip of the tail is almost entirely yellowish white. Great individual variations have been shown in this species.

Measurements: (of the type) total length 900mm; tail 290mm; foot 85mm. (Museu Paraense Emilio Goeldi) head and body 320mm; tail 410mm; foot 80mm.

Remarks: Type in the Munich Museum. Type of C. discolor in the Paris museum.

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.

Sanborn, 1949

Callicebus cupreus cupreus

Locality: Yarinacocha (1 male, 1 female, 3 skulls only, were collected at this locality for the Chicago Natural History Museum).

Sanborn, C.C. (1949). Mammals from the Rio Ucayali, Peru. Journal of Mammalogy 30(3): 284.

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.

Bender and Mettler, 1958

Callicebus cupreus

This species has 2n = 46 chromosomes, of which 10 metacentrics, 10 subterminals and 24 telocentrics. The sex chromosomes are one subterminal (X) and one telocentric (Y).

Bender, M.A. and Mettler, L.E. (1958). Chromose studies of primates. Science 128:186-190.

Cabrera, 1958

Callicebus cupreus cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Cebus cupreus (Fischer, 1829); Saguinus moloch variation (Lesson, 1840); Callithrix cupreus (Schinz, 1844); 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).

Distribution: Extreme west of Brazil, en the river-basin of the upper Solimoes and the upper Jurua, and the north of Peru east of the Andes.

Remarks: Given that the pelage of the monkeys of this genus always has shows some individual variation, it does not appear that C. ustofuscus of Elliot, based on only one specimen without locality more precise than “Brazil”, can be separated by its darker colour of the more common typical cupreus. And the paenulatus of the same author, from Andoas, the length of its pelage, by which it is distinguished, could very well be a seasonal character, or maybe a population. It is significant that Elliot includes Andoas in the localities of the real cupreus.

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).

Type locality: Forest of Rio Solimoes, Brazil. Type in Munich Museum, with three cotypes. Type locality of C. discolor Sarayacu Mission, Rio Ucayali, eastern Peru. Type in Paris Museum.

Distribution: Total range unknown; typical cupreus appears to have its centre of differentiation in the area of the Solimoes on both sides of the Peruvian-Brazilian border. The type locality is the Brazilian Solimoes, but discolor was described from eastern Peru. Lönnbergs material comprised specimens from the left bank of the Rio Jurua at Joao Pessoa, another specimen from Igarape do Gordao, also on the left bank of the Jurua and six females from the right bank of the same river and the left bank of the Rio Eiru near their confluence (at San Antonio).
Bartlett (1971) recorded cupreus at Cashiboya on the Rio Ucayali and at Santa Cruz on the Rio Huallaga. Vieira (1948) also records typical cupreus from several localities on the Upper Jurua and the Rio Eiru. Sanborn (1949) recovered typical cupreus from Yarinachocha on the Rio Ucayali.

Description: Face black, almost nude, with sparse, stiff black hairs on sides of nose and chin and shorter white hairs on lips; ear-tufts red. Forehead with narrow superciliary band of black hairs interspersed with whitish and tawny hairs; lacking contrasted light or dark frontal bar; crown speckled with strong tawny tinge, individual hairs light greyish brown at base, followed by three zones respectively yellowish-white, black and bright tawny proximo-distally, the last forming the predominant general colour; rest of upper parts, including base of tail, similar to crown, but yellowish-white most extensive on hairs, and general tone is paler on flanks and thighs than mesially, the two terminal bands being less vivid. Sides of head, chin, throat, fore-limbs (including hands), chest, belly, medial aspect of thighs, legs below knee (including feet) coppery red (mahogany-red→chesnut→maroon Lönnberg), rather darker at wrists and ankles, whit sprinkling of whitish hairs on hands and feet. Tail with rufous tinge at base dorsally, but on remainder the hairs are yellow basally, followed by brownish-black band and an apical tawny one; at end of tail the dark band is obsolete and the hairs almost wholly yellowish-white.
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.

Measurements: Males: head and body 315/350mm; tail 475/440mm; foot 95/92mm. Females: head and body 325/310mm; tail 440/420mm; foot 100/95mm.

Skull: measurements in publication

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

Hill, W.C.O. (1960). Primates. Comparative anatomy and taxonomy  4 (A): 98-147.

Hershkovitz, 1963

Callicebus moloch cupreus

Synonyms: Callithrix cuprea (Spix, 1823); Callicebus cupreus (Ihering, 1904); Callicebus cupreus cupreus (Cruz Lima, 1944); Callithrix caligata (Wagner, 1842); Callicebus caligatus (Thomas, 1908); Callicebus usto-fuscus (Elliot, 1907); Callicebus cupreus usto-fuscus (Cruz Lima, 1945); Callicebus egeria (Thomas, 1908); Callicebus cupreus egeria (Lönnberg, 1939); Callithrix discolor (I. Geoffroy, 1851 –part: not 1848, 1852).

Type locality: Rio Solimoes, Amazonas, Brazil, near the Peruvian boundary, here restricted to Tabatinga. Cotypes in Münich and Leiden Museums.

Distribution: South of the Amazon from the left bank of the Rio Madeira, Amazonas, Brazil, to the Rio Yavari and mouth of the Ucayali in Loreto, Peru; altitudinal range approximately 200 meters asl.

Description: (key to species:) general body colour grey, reddish or brown; under parts like back or sharply defined reddish orange or buff; hind feet black, brown, red or grey, tail grey or blackish with tip grey or grey mixed with black; throat like chest; forearms grey, red, dark brown sometimes blackish above; upper surface of hands grey to blackish never sharply contrasted with colour of upper side of wrists.
Forehead grey sometimes edged with black or dominantly black to blackish brown and always well defined from posterior half of crown and nape, outer side of forearms grizzled or more or less uniformly reddish to dark brown. Forehead blackish; under parts of body reddish brown or blackish brown and not always well defined from sides; forearms chestnut, reddish brown or blackish; tail grey, dark brown or blackish. Under parts and forearms reddish an usually well defined from sides; forearms chestnut, reddish brown or blackish; tail grey, dark brown or blackish.

Measurements: see table in publication.

Remarks: The specimens in the Chicago Natural History Museum from Tabatinga, the Rio Yavari and the Rio Maniti, all in the type region of cupreus, agree completely with the description and coloured plate of the original specimen. In these the crown is reddish brown but with the black of the forehead extending over the anterior portion. The hands and feet are reddish more or less washed with black.
Lower down the Solimoes, grey-crowned titis appear which are otherwise indistinguishable from topotypical cupreus. These suggest an intimate relationship with and the postulated derivation from, discolor. Callicebus egeria is based on a specimen from Ega (= Tefé) was distinguished from cupreus by its greyish crown. Later Thomas (1928) recorded a series of cupreus “collected by Ehrhardt in the typical region of the Solimoes which shows that the colour of the crown is so variable – rufous or grey that C. egeria should probably be united with C. cupreus”. Lönnberg (1939) recorded 2 individuals from Jaburú, Rio Purús, as similar to egeria in the colour of their crown. He also noted other specimens with typical cupreus type heads from localities higher up, lower down and on both sides of the river. These he assigned to Callicebus caligatus.
A specimen from Lago da Ayapuá, left bank of the lower Rio Purús differs from typical cupreus by the presence of a grey frontal blaze followed by a dark band and the reddish brown crown usual in cupreus. The colour pattern of the head of the Brazilian specimen conforms to that of C. moloch ornatus. Pelzeln (1883) had previously recorded a similar individual of unknown origin, purchased in 1840 from a dealer. He described it as indistinguishable from cupreus except for the ornatus-type colour pattern of its head.
It seems that there is a considerable amount of variation in the head pattern with a tendency for the grey crown or forehead to dominate in the area between the lower Purús and Juruá. Higher up the Solimoes along the Maranon and Ucayali, black fronted cupreus grades info white fronted discolor.
The original description of Callithrix caligatus agrees precisely with that of cupreus. Said to be from Borba, Rio Madeira and from Manaqueri, Rio Solimoes, its type locality was since restricted to the former locality by Thomas. The town of Borba, however, is on the right side of the Madeira, where only hoffmannsi occurs. It seems necessary to assume, therefore, that the type of caligata originated west of the Madeira, probably in the forests opposite the town of Borba. Its cotype from Manaqueri and all other recorded specimens of cupreus are from west of the Madeira.
The type of Callicebus ustofuscus from “Brazil” was collected by Castelnau and its description applies only to cupreus. The individual depicted in vivid reds by Cruz Lima as representative of ustofuscus is also undoubted cupreus. Hill (1960) describes under the name Callicebus cupreus ustofuscus an animal having “forehead with black band succeeded by whitish bar as in leucometopus (= discolor)”. On the facing page he presents a photograph captioned Callicebus cupreus of a different titi with broad black frontal band extending onto the crown. Hill notes that the type of ustofuscus is in the British Museum but his account of the animal is ostensibly derived from Cruz Lima who only speculated on the true nature of a specimen he never saw. Plates 1 and 2 poses of the same individual figured by Hill as ustofuscus. The animal lived in 1952 in the zoological park of the New York Zoological Society. Its colour pattern is that of cupreus but the precise colouration cannot be determined from the photograph. The hands appear to be grey while in true cupreus they are reddish.
Callicebus moloch cupreus as now constituted may prove to be composite. It is certainly polyphyletic as attested by characters which point to a brunneus stock ancestry of some populations and a discolor stock ancestry of others. Although separated from the range of hoffmannsi by breadth of the Rio Madeira only, its relationship to that form must be traced through the donacophilus-brunneus chain.

Specimens examinedBrazil – Amazonas: Lago do Ayapuá. Peru – Loreto: Quebrada Esperanza, Rio Yavari Mirim; Santa Cecilia, Rio Maniti; San Fernando, Rio Yavarí; Tbatinga, Rio Maranon.


Egozcue, et al., 1969

C. moloch (according to Hershkovitz 1990 this is a C. cupreus!)

Diploid number 2n = 46; 10 pairs of submetacentrics, 12 pairs of acrocentrics. X is submetacentric and Y is a minute element that can be identified as submetacentric.

Egozcue, J., Perkins, E.M., Hagemenas, F. and Ford, D.M. (1969). The chromosomes of some Platyrrhini (Callicebus, Ateles and Saimiri). Folia primatologia 11: 17-27.

De Boer, 1974

Callicebus moloch cupreus

2 individuals, originating from Peru.
Diploid number 2n = 46; 24 biarmed and 24 acrocentric chromosomes.

The karyotype described is identical to those reported for C. cupreus by Bender and Mettler (1958), who identified an only slightly different X, and for C. moloch (=cupreus) by Egozcue et al. (1969).

The arrangement and the structure of the autosome pairs are nearly identical to those of C. moloch ornatus.

The cytological information suggests that the forms cupreus and ornatus on the one hand, and donacophilus on the other, may be more than only subspecifically distinct.

Boer, L.E.M. de (1974). Cytotaxonomy of Platyrrhini. Genen Phaenen 17: 28-37.

Freese, 1982

Callicebus moloch

Locality: Orosa (3º22’S-72º07’W to 3º46’S-72º20’W).

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.

Kraft, 1983

Callicebus cupreus cupreus

Synonym: Callithrix cuprea

Remark: Four type specimens present in Munich collection.

Kraft, R. (1983). Die von JB Spix beschriebenen neotropischen Primaten und Chiropteren Verzeichnis der in der Zoologischen Staatssammlung München. Spixiana Supplement . 9:429-441.

Hershkovitz, 1987

Callicebus cupreus cupreus

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

Hershkovitz, 1988

Callicebus cupreus cupreus

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. Juvenals 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.

Spread of the prototypical C. cupreus to its present eastern boundary along the west bank of the Río Purús is evidently recent. Differentiation of the populations between the Río Juruá and Purús is slight, but additional material, not now available for comparison, may prove the difference significant. The name Callicebus cupreus egeria is available for the titis between the Ríos Purús and Juruá.

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.

Peres, 1988

Callicebus moloch cupreus

Localities: Brazil, Acre, Sao Domingos (between upper Rio Purus and Rio laco; 8055’S, 68°20’W); Brazil, Amazonas, Igarape-Açu (left bank of lower Rio Urucu (4°30’S, 64°29’W); Brazil, Amazonas, Lago da Fortuna (left bank of Rio Jurua; 5°05’S, 67°10’W); Brazil, Amazonas, SM-1 (oil-drilling site between Rio Urucu and Rio Coari; 4°50’S, 65°1 6’W).

Peres, C. A. 1988. Primate community structure in western Brazilian Amazonia. Primate Conservation 9: 83-87.

Soini et al., 1989

Callicebus moloch cupreus

Distribution: Peru.


Hershkovitz, 1990

Callicebus cupreus cupreus

Member of th moloch Group.

Synonyms: Callithrix cuprea (Spix, 1823); Callicebus egeria (Thomas, 1908); Callicebus toppini (Thomas, 1914); Callicebus acreanus (Vieira, 1952); Callithrix discolor (I. Geoffroy, 1851 – part).

Type locality: Rio Solimoes, Brazil, near the Peruvian boundary; restricted to Tabatinga by Hershkovitz (1963a, p. 36), but should be Rio Solimoes opposite Tabatinga because the species does not occur on the north bank or Tabatinga side of the Solimoes. Restriction of the type locality of Callicebus cupreus Spix to the “Peruvian Amazonas” by Thomas (1908, p. 90) is not valid. The types originated in Brazil. Lectotype in the Zoologische Staatssammlung, München.

Distribution: South bank of the Amazonas-Solimoes from the left bank of the Rio Purus, Amazonas, Brazil, west to the east bank of the Rio Ucayali, in Loreto and northern Ucayali, Peru, south in the Rio Purus basin in Acre, Brazil, Loreto, and Madre de Dios, Peru; altitudinal range to approximately 200 m above sea level. Specimens in the American Museum of Natural History from Rio Urubamba and from Sarayacu may have been collected elsewhere.

Description: Sideburns, legs, and under parts uniformly reddish contrasting with buffy agouti of upper and outer sides of trunk and crown; forehead like crown but usually with blackish fringe formed by superciliary vibrissae and bases of marginal hairs.

Measurements: See publication.

Comparisons: Distinguished from Callicebus cupreus ornatus and nearly all C. c. discolor by absence of contrastingly pale transverse frontal blaze; however, a few dominantly whitish frontal hairs sometimes present; from C. c. ornatus and C. oenanthe by uniformly reddish arms and cheiridia; from C. caligatus and C. brunneus by entire crown buffy to orange agouti, cheiridia reddish; from all other forms of Callicebus by one or more of the above characters.

Specimens Examined: Total 130. Brazil – Acre: Iquiri (holotype of acrensis); Manoel Urbano; Sao Luiz da Mamoria; Sena Madureira; Amazonas: Aiapua; Eirunepe; Fonte Boa; Igarape Gordao; Itaboca; Jaburu; Joao Pessoa; Pauini; Porta da Castanha, Tefe; Rio Jurua; Santa Cruz; Santo Antonio; Tefe (holotype of egeria). Peru – Loreto: Balta; Cashiboya; Cerro Azul; Orosa; Pavas; Madre de Dios: Rio Inuya; Rio Tapiche; Rio Yavari; “Sarayacu,”; Tahuamanu (holotype of toppini); Ucayali: “Rio Urubamba”.

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

Puertas and Bodmer, 1993

Callicebus cupreus

Locality: Tamshiyacu-Tahuayo Comunal Reserve; situated in north-eastern Peru between the Tamshiyacu, Tahuayo and Yavari Miri rivers.


Schneider, et al., 1993

Callicebus cupreus

Locality: seized at the margins of the Maniti River, right bank of the Amazon, Peru.

Remarks: Estimates of genetic similarity between the taxa showed that C. moloch is the most differentiated, since it presented genetic distances of 0.059 and 0.066 when compared to C. brunneus and C. cupreus, respectively; the most distinctive loci are GPI and CA2. On the other hand, the genetic distance between C. brunneus and C. cupreus showed a value of 0.008 only. This estimate is quite similar to those observed among other New World monkeys at the subspecies level. In fact, assuming Thorpe’s (1982) criteria, that genetic distances lower than 15% would be typical of those separating subspecies, we would have to consider the three taxa investigated here as belonging to a single species. As shown in the table, cytogenetic studies in the genus Callicebus are scarce. The karyotypes of eight of the 13 species in this genus are still unknown. However, the three species considered in the present study have already been karyotyped. Consideration of the biochemical and cytogenetic data together, therefore, establishes that:
a) the comparison which showed the largest chromosome difference (brunneus-cupreus) is the one presenting the least differentiation at the biochemical level; and
b) the cytogenetic diversity is compatible with divergence at the species level, unlike the biochemical data.

Schneider, H; Schneider, M.P.C.; Sampaio, M.I.C.; Montoya, E.; Tapia, J. ; Encarnacion, N.P. ; Anselmo, N.P. and Salzano, F.M. (1993). Divergence Between Biochemical and Cytogenetic differences in three Species of the Callicebus moloch Group. American Journal of Physical Anthropology 90:345-350.

Peres, 1993

Callicebus cupreus cupreus

Locality: both sides of the Juruá River, Western Brazilian Amazonia.

Remarks: collected specimens deposited in the Museu Goeldi, Belém, Brazil.

Peres, C.A. (1993). Notes on the primates of the Juruá river, western Brazilian Amazon. Folia Primatologica 61:97-103.

Aquino and Encarnacion, 1994

Callicebus cupreus cupreus

Distribution: C. c. cupreus is distributed from the Amazon River in the north to the Rio Purus in the Department of Ucayali in the south, and from the Rio Ucayali in the west to the Brazilian border in the east.

Description: Pelage generally greyish-brown. Forehead and crown dark-brown, reddish or grey. Upperparts, upper arm and thigh greyish-brown, forearm and lower leg reddish-brown to light-reddish. Under parts and inner sides of limbs varying from reddish-brown and reddish-orange to buffy. Tail grey or blackish, terminal portion hoary. In C. c. cupreus the forehead is dark-brown.

Measurements: Total length up to 85cm and weight of 1200-1500g.


Kobayashi, 1995

Callicebus cupreus cupreus

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.

 Kobayashi, S. (1995). A phylogenetic study of Titi Monkeys, Genus Callicebus, based on cranial measurements: 1. Phyletic groups of Callicebus. Primates 36(1): 101-120.

Voss and Emmons, 1996

Callicebus cupreus

Localities: Brazil, Acre, Sao Domingos (between upper Rio Purus and Rio Iaco; 8055’S, 68°20’W); Brazil, Amazonas, Igarape-Açu (left bank of lower Rio Urucu (4°30’S, 64°29’W); Brazil, Amazonas, Lago da Fortuna (left bank of Rio Jurua; 5°05’S, 67°10’W); Brazil, Amazonas, SM-1 (oil-drilling site between Rio Urucu and Rio Coari; 4°50’S, 65°1 6’W); Peru, the Cashinahua indian village of Balta (10°08’S, 71°13’W; ca. 300 m elevation), in Departamento Ucayali (formerly Loreto); Peru, Loreto, Rio Orosa (average coordinates of survey ca. 3°34’S, 72°14’W).

Remarks: Five specimens of titis from “Orosa” in the AMNH were listed by Hershkovitz (1990: 62) as examples of Callicebus cupreus and six others as C. caligatus (op. cit.: 66). However, the AMNH has only six specimens of Callicebus from Orosa, not eleven; the skin characters of this series (AMNH 73703-73708) match those described (op. cit.: 61) for C. cupreus cupreus. The report of two sympatric members of the moloch group of titis at Orosa was an error caused by inadvertently listing both original and revised identifications of the same series among the specimens examined (Hershkovitz, in litt.). Thus, Callicebus cupreus is presumably the correct identification for the monkeys Freese et al. (1982) reported as C. moloch.

Voss, R.S. and Emmons, L.H. (1996). Mammalian diversity in neotropical lowland rainforests: a preliminary assessment. Bulletin of the American Museum of Natural History 230: 1-114.

Peres, 1997

Callicebus cupreus

Localities: both sides along the Rio Juruá, Brazil.

Peres, C.A. (1997). Primate community structure at twenty western Amazonian flooded and unflooded forests. Journal of Tropical Ecology 13:381-405.

Aquino, 1998 

Callicebus cupreus

Localities: Quebrada Blanco, an affluent of the Río Tahuayo, sout-east of Iquitos (4°23’S , 72°55’W), more precisely the area delimited by Quebradas Tunchío, Palmichal and Tangarana. Río Yavarí, to the east of Iquitos, between Agua Negra and Carolina (4°30’S , 71°43’W) (callicebus.nl: it is not clear if Aquino reports that the species occurs at both localities, as he also mentions C. caligatus).

Aquino, R. (1998). Some observations on the ecology of Cacajao calvus ucayalii in the Peruvian Amazon. Primate Conservation 18: 21-24.

Calouro, 1999

Callicebus cupreus cupreus

Localities: Parque Nacional da Serra do Divisor, near the border with Peru . Sítio Norte 5: 07°27’32″S, 73°46’28″W; Sítio Norte 9: 07°27’02″S , 73°36’30″W; Sítio Norte 6: 07°21 ’23″S, 73°40’41″W; Sítio Norte 8: 07°33’24″S, 73°16’36″W; Sítio Sul 1: 08°16’51″S, 73°15’13″W.


Bennett et al., 2001

Callicebus cupreus

Localities: seasonally flooded forest along the Tapiche River in north-eastern Peru at 5°39’18’’ S by 74°00’13’’ W with an elevation of approximately 110 m.


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).

Distribution: South of the Napo-Solimoes, from the Rio Purus-Ituxi to the Andes; Hernandez-Camacho and Cooper (1976) mapped it as far north as the southern bank of the Rio Guamues, in extreme southern Colombia. A population on the Rio Sucusari, a lower left-bank tributary of Rio Napo, was reported by Brooks and Pando-Vasquez (1997).

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. The distributions of the four, taken as a group, are coterminous, but the two easterly forms have the frontal blaze whereas the two westerly ones are without it, so that at the very least there is a remarkable parallelism between phaeomelanic and eumalenic species in their geographical variation. I suggested (Groves, 1992) that this odd situation could be resolved if the eumelanic and phaeomelanic “species” are actually morphs of a singe species.
This hypothesis was tested by my examination of the skins in AMNH in 1997. It is corroborated; not only are eu- and phaeomelanic skins most readily interpreted as morphs of a single species, there is in fact very little difference between them at all. In skins identified as caligatus, the hair bases are redder, less brown than those labelled cupreus; the underside is more maroon, less foxy red; the red of the hands and feet is darker; the maroon of the cheeks is blackish rather than red.
Close reading of Hershkovitz (1990), indeed, suggests that even the geographic variation in presence or absence of the frontal blaze may not be so clear-cut. Thus, two series ascribed to C. c. discolor, collected by the Olallas at the mouth of the Rio Inuya (Rio Urumbamba) and Lagarto (upper Ucayali), were regarded as misplaced by Hershkovitz because they are on the “wrong” side of the Ucayali. He similarly regarded the localities of Ollala specimens of C. caligatus (Sarayacu and mouth of Rio Inuya). If the localities of these specimens are indeed incorrect, then an easterly trend in development of the frontal blaze is very marked; if not (if the Olallas correctly reported the localities), then blaze development ceases to have much geographic significance.
Finally, Hershkovitz (1990) also queried the locality of the type specimen of his C. dubius. If it originated from the east bank of the Rio Purus, as he hypothesized, then C. dubius is preserved as a distinct taxon with a small, river-bounded range; if from the west bank, as the label records, then the taxon has no identifiable separate distribution. From my findings and conclusions given here, I see no reason at all to query the locality as given.


Heymann et al., 2002

Callicebus cupreus

Localities: 2º09’S, 75º00’W Quebrada Arabela; 2º07’S, 74º52’W Buena Vista; 2º03’S, 74º54’W (Playa); 2º10’S, 74º39’W (Casa Calvo); 2º16’S, 74º28’W Soledad; 2º15’S, 74º28’W Soledad.


Roosmalen et al., 2002

Callicebus cupreus

Type locality: Right bank of Rio Solimões, Brazil, near the Peruvian border. The lectotype is an adult female, mounted, including skull, Zoologische Staatssammlung, München, Germany, no. 10; lectoparatypes in the same collection are no. 24, no. 89 a + b, all collected January, 1820 by J. B. von Spix.

Description: Sideburns, sides of neck, throat, inner surface of limbs, and under parts of body uniformly reddish, sharply contrasting with buff-brown agouti of dorsum and outer sides of trunk, basal part of tail, and crown; forehead as crown, reddish-brown agouti, often fringed with blackish superciliary vibrissae and marginal hair bases.
Distinguished from other members of the cupreus group, except for C. caligatus and C. stephennashi, by absence of distinct pale transverse frontal blaze, and from all members
of the cupreus group by an overall dark tail with pencil only white; from Callicebus caligatus by lacking a broad black frontal transverse blaze; from Callicebus stephennashi by lacking a black frontal transverse blaze that contrasts sharply against a silvery crown, and having an overall dark tail instead of a three-quarters to entirely buffy to white tail; from Callicebus brunneus by its red sideburns, forearms, lower legs, cheiridia and under parts of the body.

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.

Oliveira Azevedo Lopes and Rehg, 2003

Callicebus cupreus

Locality: Serra do Divisor National park, the western edge of Acre, Brazil.


Tirado Herrara et al., 2003

Callicebus cupreus

Locality: Estación Biológica Quebrada Blanco, northeasten Peru (4º21’S, 73º09’W).

Tirado Herrera, E.R.; Franke, T.; Knogge, C. and Heymann, E.W. (2003). Flower and Fruit Visitors of Marcgravia longifolia in Amazonian Peru. Plant Biology 5(2):210 – 214.

Dumas et al., 2005

Callicebus cupreus

The karyotype has a diploid number of 2n=46 and autosomal FN=33 confirming previous results (de Boer 1974; Benirschke and Bogart, 1976). We found eleven pairs of meta/submetacentric autosomes and eleven pairs of acrocentric ones. The sex-chromosome system is XX/XY. The X chromosome is a submetacentric, typical for most mammals and Y is a very small acrocentric

Blood samples of 79 Callicebus cupreus individuals were kindly provided by California Regional Primate Research Center, University of California, Davis. The founders of the colony were obtained in the early 1970s from Iquitos (Peru). Other animals from the same location were added in the early 1990s.

Remarks: No differences in G-banding pattern were noted in the 79 individuals studied; therefore the two groups of monkeys that formed the colony between the early 1970s and the 1990s must have had the same karyotype.

Dumas, F.; Bigoni, F.; Stone, G.; Sineo, L. and Stanyon, R. (2005). Mapping genomic rearrangements in titi monkeys by chromosome flow sorting and multidirectional in-situ hybridization. Chromosome Research 13: 85–96.

Bicca-Marques, 2005

Callicebus cupreus cupreus

Locality: Zoobotanical Park (9°56’ 30”–9°57’ 19” S, 67°52’ 08”–67°53’ 00” W) of the Federal University of Acre located in Rio Branco, State of Acre, Brazil.


Rehg, 2006

Callicebus cupreus

Locality: Fazenda Experimental Catuaba (S10°04’S,067°36’W) is ca. 25 km east of the Rio Acre (Acre River) and Rio Branco, the capital city of the state of Acre in north-western Brazil.

Rehg, J.A. (2006). Seasonal Variation in Polyspecific Associations Among Callimico goeldii, Saguinus labiatus, and S. fuscicollis in Acre, Brazil. International Journal of Primatology.  27(5): 1399-1428.

Nadjafzadeh and Heymann, 2008

Callicebus cupreus

Locality: Estación Biológica Quebrada Blanco (4º21’S, 73º09’W) altitude of about 120 m a.s.l., on the right bank of Quebrada Blanco, an affluent of the Rio Tahuayo, in north-eastern Peru.

Nadjafzadeh, M.N. and Heymann, E. W. (2008). Prey Foraging of Red Titi Monkeys, Callicebus cupreus, in comparison to sympatric tamarins, Saguinus mystax and Saguinus fuscicollis.  American Journal of Physical Anthropology 135:56–63.

Haugaasen and Peres, 2009

Callicebus cupreus

Locality: Lago Uauaçu in the lower Rio Purús region of central-western Brazilian Amazonia.

Haugaasen, T. and Peres, C.A. (2009). Interspecific primate associations in Amazonian flooded and unflooded forests. Primates 50: 239–251.

Vermeer, 2009

The diagnostic characters of Callicebus cupreus are described by Van Roosmalen et al. (2002), and depicted in a drawing by Stephen Nash. The description and the drawing were compared to the lectotypes and the lectoparatypes of Callicebus cupreus at the Zoologische Staatssammlung in München (Nos. 10, 24, 89a and 89b). The most important difference between the drawing in the publication and the lectotype is the color of the tail (the color of the tail is not described by Van Roosmalen et al., 2002). While the tail of the animal in the drawing is the same buff-brown agouti color as its hindlimbs, the tail of lectotype No. 10 is much lighter, comparable to that on the drawing of Callicebus moloch in the publication of Van Roosmalen et al. (2002). The tail of lectoparatype No. 24 is identical to that of the lectotype, while the tails of the paralectotypes 89a and 89b are somewhat darker. Most other specimens of Callicebus cupreus that I have examined in the collections of the American Museum of Natural History in New York and the Naturalis Museum in Leiden have lighter and more greyish colored tails than the ones depicted by Van Roosmalen and colleagues (2002). Observations in the wild by Eckhard Heymann, at Estación Biológica Quebrada Blanco (4° 21′ S, 37° 09′ W), well within the known distribution of Callicebus cupreus, confirm that the tail of adult Callicebus cupreus is greyishwhite (Eckhard Heymann, pers. comm.). The tail of young Callicebus cupreus is brownish, but has the greyish colour of the adults by approximately 2 years of age (pers. obs. at La Vallée des Singes, Romagne, France). The captive population in European zoos is partly based on individuals that were captured near the Rio Maniti in Peru by the California National Primate Research Center of Davis. Rio Maniti is also within the distribution of Callicebus cupreus. All these animals have greyish tails, strikingly different than the colour of their back and legs.

Vermeer, J. (2009). On the Identification of Callicebus cupreus and Callicebus brunneus. Neotropical Primates 16(2): 69-71.

Auricchio, 2010

Callicebus cupreus

Localities: see publication.

Description:
Phenotype 1: Face reddish-cream; Forehead, crown, nape and back reddish-cream (agouti hair banded with light stripes longer than dark ones); Lower back reddish-cream (agouti hair banded with light stripes longer than dark ones), but washed with brown; External surface of fore legs and forearms intense redish brown which can vary to orangish; Back of hands, fingers and back of feet brown, not agouti; Base of tail reddish-cream (agouti hair banded with light stripes longer than dark ones), but washed with brown; Middle tail reddish-cream (agouti hair banded with light stripes longer than dark ones), but washed with brown; Tip of tail reddish-cream (agouti hair banded with light stripes longer than dark ones), but washed with brown; Ventral surface intense reddish brown which can vary to orangish.

Phenotype 2: Two specimens from Pauini have arms, legs, chest and ventral surfaces orangish.

Phenotype 3: A specimen from the Iquiri River, the holotype of C. cupreus acreanus and two specimens from Santa Cruz do Eiru River have the forehead and crown agouti-brown with black and cream, lighter than described for phenotype 1, back as moloch and lower-back more brownish. Tail is dark-brown, gradually getting lighter to the tip, which is cream. Arms, legs, ventral surfaces and beard are dark reddish-brown, almost dark red.

Six specimens (MPEG 1587, 1588, 1605, 1608, 1609 and 1845) from Amazonas (Rio Javari, Estirão do Equador) are darker than the phenotype 3, described here. The phenotypes are distributed in four localities that are inside the known distribution of C. cupreus and do not show a geographic pattern that could suggest an existence of more than one only taxon. As it was not possible to identify geographical limits that could indicate segregation among taxa, and it was not possible to perform a DNA analysis, definite considerations about the taxonomic status of C. cupreus must await, intra-specific color polymorphism being the best explanation for the observed pattern.

Remarks: Groves (2001) follows Hershkovitz (1990) in Callicebus taxonomy, but doubts him concerning some propositions. One of them considers C. caligatus, C. dubius and C. cupreus as synonyms. Roosmalen et al. (2002) described differences among these three species, considering all of them valid, a view I agree based on morphological grounds. All three show several distinctive characters, as pointed out by Roosmalen et al. (2002) and revised here, such as the presence or absence of chromogenetic fields, e.g. frontal white and black stripes, tip of tail and white fingers.

Auricchio, P. (2010). A morphological analysis of some species of Callicebus. Neotropical Primates 17(2): 47-58.