Literature paenulatus

Thomas, 1880

Callithrix cuprea

Locality: Copataza River, Ecuador. Andoas, Ecuador.

Remarks: The Andoas specimen, which is a male, differs from the rest in having the fur of the back of a dirty orange-grey colour, without annulations, instead of being a bright annulated black and white. One of the others, a female, shows a tendency to this condition of the hair, which is therefore very probably a seasonal change, as the Andoas specimen was shot in September, while the others were obtained between December and February.

Thomas, O. (1880). On mammals from Ecuador.  Proceedings of the Scientific Meetings of the Zoological Society of London.  Pp. 394.

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”.

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

 Elliot, 1909

Callicebus paenulatus

Type locality: Andoas, on the Pastasa River, Ecuador.

Description: Fur very long, thick, and fluffy; mantle uniform colour, distinct from back; arms from elbows hands, feet, and legs above knees uniform colour. Forehead grizzled ochraceous, the hairs black, with ochraceous tips, this being the dominant colour; crown and occiput rich cinnamon-rufous, this colour produced by the tips of the hairs; mantle uniform tawny ochraceous, extending behind shoulders; middle of back dark hair-brown, grading into tawny ochraceous on the rumps; flanks dark grizzled brownish grey; hairs on ears, cheeks, arms from elbows, hands, legs from above knees, feet, inner side of limbs, chin, throat, and under parts of body rich dark burnt sienna; tail very long, grizzled white and black, the orange-buff of the basal portion of the hairs showing through, darkest at base, and the tip buff; the rest of the tail is tawny ochraceous like the rump; flanks dark grizzled brownish grey; hairs on ears, cheeks, arms from elbows, hands, legs from above knees, feet, inner side of limbs, chin, throat, and under parts of body rich dark burnt sienna; tail very long, grizzled white and black, the orange-buff of the basal portion of the hairs showing through, darkest at base, and the tip buff; the rest of the tail is tawny ochraceous like the rump.

Measurements: Total length 850mm; tail 520 (skin).

Skull: occipito-nasal length 29.1; zygomatic width 40.6; palatal length 29.1; length of nasals 9.3; length of upper molar series 14.2; length of mandible 41.3; length of lower molar series 15.8.

Remarks: Type an adult, in British Museum, no. 80.5.6.14.
This species differs in many ways from all of the others in the genus. It is the only one of the cuprea style with a uniformly coloured mantle and one so long and thick as to resemble a mane. The red on arms and legs extends much higher than in any other species. A single specimen was procured by Mr. Buckley at Andoas.

Elliot, D.G. (1909). Descriptions of apparently new species and subspecies of monkeys of the genera Callicebus, Lagothrix, Papio, Pithecus, Cercopithecus, Erythrocebus and Presbytis. Annals and Magazine of Natural History 8 (4): 244-245.

Elliot, 1913

Callicebus paenulatus

Synonyms: Callicebus paenulatus (Elliot, 1909).

Type locality: Andoas on the Pastas River, Ecuador (British Museum).

Description: Fur very long, thick and fluffy; mantle uniform colour, distinct from back; arms from elbows, and hands, feet and legs to above knees, uniform colour. Forehead grizzled ochraceous, the hairs black with ochraceous tips, this being the dominant colour; crown and occiput rich cinnamon rufous, the colour produced by tips of hairs; mantle tawny ochraceous uniform, extending behind shoulders; middle of back dark hair brown, grading into tawny ochraceous on the rump; flanks dark grizzled brownish grey; hair on ears, cheeks, arms from above elbows, hands, legs from above knees, feet, inner side of limbs, chin, throat and under parts, rich dark burnt sienna; tail very long, grizzled white and black, the orange buff of the basal portion of the hairs showing through, darkest at base and the tip buff. The root of the tail is tawny ochraceous like the rump.

Measurements: Total length, 850mm; tail, 520mm (skin).

Skull: occipito-nasal length, 57.3; Hensel, (skull broken); zygomatic width, 40.6; palatal length, 29.1 ; median length of nasals, 9.3; length of upper molar series, 14.2; length of mandible, 41.3; length of lower molar series, 15.8.

Remarks: This species differs in many ways from all others of the genus. It is the only one of the C. cupreus style with a uniformly coloured mantle, and one so long and thick as to resemble a mane; the red on arms and legs extends much higher than on any other species, and the fur everywhere is longer and thicker. A single specimen was procured by Mr. Buckley at Andoas.

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

Lönnberg, 1922

C. paenulatus (Elliot) appears also to belong to the same group, but, although it is similar with regard to the colour of the lower parts and legs, it seems to be very different in having a “mantle”, which is said to be ‘tawny ochraceous uniform, extending behind the shoulders; middle of the back dark hair brown, grading into tawny ochraceous on the rump” (Elliot); and Thomas speaks about an “elongated mantle, paler than the rest of the back” in the same species. This is, of course, entirely different from the condition of the present specimens (= C. cupreus napoleon), in with the hairs are nearly of equal size all over the upper parts, about 40mm., or a little more, perhaps partly 45mm. on the neck. Likewise the hairs are ringed everywhere on the body, so that there is no trace of a pale and unicoloured mantle.


Thomas, 1928a

Remark: C. paenulatus (Elliott) is probably the same as C. cupreus.


Cruz Lima, 1945

Callicebus cupreus paenulatus

Synonym: Callicebus paenulatus (Elliot, 1909).

Description: Forehead grizzled ochraceous, the hairs black, with ochraceous tips, this being the dominant colour; crown and occiput rich cinnamon-rufous, this colour produced by the tips of the hairs; mantle uniform tawny ochraceous, extending behind shoulders; middle of back dark hair-brown, grading into tawny ochraceous on the rumps; flanks dark grizzled brownish grey; hairs on ears, cheeks, arms from elbows, hands, legs from above knees, feet, inner side of limbs, chin, throat, and under parts of body rich dark burnt sienna; tail very long, grizzled white and black, the orange-buff of the basal portion of the hairs showing through, darkest at base, and the tip buff; the rest of the tail is tawny ochraceous like the rump. (Original description)

Measurements: (skin), total length 850mm; tail 520mm.

Remarks: Thomas in 1880 referred to this animal as C. cupreus, attributing the difference in colouration to a possible seasonal variation, adding further that one female specimen from the Copataza in a series of twelve of C. cupreus showed a tendency toward these conditions.
In some specimens of forms from the lower Amazon we had the opportunity of observing also a pronounced tendency toward unicolouration of the hairs on the dorsal surface which, to a certain extent, led to our doubting of the validity of the species C. paenulatus. The author points out other distinguishing features based on the fur, as, for instance, its length and thickness, which, however, are not sufficient to allay our doubts. Only with the acquisition of a greater number of specimens for comparison and the verification of the assumption of seasonal variation can the validity of such characteristics be ascertained.

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

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).

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 paenulatus

Synonyms: Callicebus paenulatus (Elliot, 1909).

Type locality: Andoas, 70 miles below Sarayacu, Rio Pastaza, Oriente, S.E. Ecuador. Type in British Museum (no. 80.5.6.14), collected by C. Buckley.

Description: Forehead grizzled ochraceous, without white band, the hairs black basally with ochraceous tips; crown and occiput rich cinnamon-rufous, the hairs tipped with this colour but basally grey; upper back uniformly tawny-ochraceous to behind shoulder, posterior to this darker – dark hair-brown grading into tawny-ochraceous on rump; flanks dark grizzled brownish-grey; hairs on ears, cheeks, arms beyond elbows, hands and fingers, legs from above knees, feet and toes, medial aspects of limbs, chin, throat and remainder of under parts rich dark burnt sienna; tail grizzled black and whitish, with some orange-buff of bases of hairs showing through; base of tail darker, tip lighter (buffy).

Remarks: the type was originally described as C. cupreus by Thomas (1880), who noted a colour difference which he attributed to seasonal variation, in view of the fact that a single female of cupreus from the Rio Copataza in a series of 12 of typical cupreus showed a tendency towards the same appearance. Lima has also shown the tendency to unifor;ity of colouration in dorsal hairs to occur in material from the Lower Amazon and therefore doubts the validity of paenulatus.

Distribution: Known only from the type locality.

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

Hershkovitz, 1963

Callicebus moloch discolor

Synonyms: 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.

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 with grey band often thinly edged with black; under parts of body orange or reddish and always well defined from sides; forearms uniformly orange or reddish or contrasting grizzled on outer side; tail dominantly grey. Forearms and hind legs uniformly rufous above and below.

Measurements: see table in publication.

Remarks: There is no reason to believe that paenulatus, for al the peculiarities ascribed to it, is taxonomically distinct from any other titi of eastern Ecuador.


Hershkovitz, 1990

Callicebus cupreus discolor

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).

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

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).