Synonymy

1766 Simia pithecia Linnaeus, Systema Naturae 12th ed. 40. Location unknown.

1789 L' Yarke, Buffon. Histoire Naturelle Generale et Particuliere, a l'Histoire des Animaux Quadrupedes, Suppl. Tome Septième, Paris: L'Imprimerie Royale. Location unknown.

1797 Simia leucocephala Audebert, Histoire Naturelle des Singes et Makis, 6^eme fam., 1: 9, plate 2. French Guiana.

1803 Callithrix pithecia (Le saki ventre roux) É. Geoffroy Saint-Hilaire. Catalogue des Mammifères du Museum National d'Histoire Naturelle, Paris, No. 9. Type no longer exists, female. French Guiana, (see “Discussion”).

1812 Simia rufiventer / Pithecia rufiventer É. Geoffroy Saint-Hilaire. Recueil d'Observations de Zoologie et d'Anatomie Comparee (Humboldt expedition), vol. 2, p.358. Type no longer exists. Female. French Guiana. Nomen oblitum. (see “Discussion”).

1818 Simia adusta Olfers. Journal von Brasilien, 2: 198. Location unknown. Hershkovitz (1987) substitute name for Simia pithecia Linnaeus.

1818 Simia nocturna Olfers. Journal von Brasilien, 2: 198. No. 39097 at RNH Leiden, subadult male, French Guiana, Cayenne.

1819 Pithecia saki Muirhead. Edinburgh Encyclopedia 13: 400. British Guiana.

1820 Pithecia rufibarbata Kuhl. Beitrage zur Zoologie und Vergleichenden Anatomie, p.44. Hershkovitz (1987) notes “holotype juvenile mounted in Leiden,” juvenile female. Suriname.

1820 Pithecia ochrocephala Kühl. Beitrage zur Zoologie und Vergleichenden A natomie, p.44. based on the same specimen as Simia nocturna Olfers, 1818, from the Temminck Voyage, subadult male. French Guiana, Cayenne.

1987 Pithecia pithecia pithecia Hershkovitz. Am. J. Primatol. 12: 418–419; in part.

Holotype. A type specimen is not known to exist (Hershkovitz, 1987).

Neotype. Simia leucocephala mount No. 452, placed in the Museum d'Histoire Naturelle, Paris in 1822 (collector appears to be S. Foiteau) is here designated as a neo type.

Type locality. Cayenne, French Guiana, for the neotype leucocephala.

Specimens examined. Two hundred and nine skins and skulls, and dozens of live, captive and wild photos.

Description. Males. Fully mature adult males are black with little or no stippling of the dorsal hair. The hands and feet are generally black with no or little hair covering them. Male facial disks are white “half moons” completely covering the circumference of the face, connected in most cases in the midline of the face by a thin black line extending from the top of the forehead to the mid-glabella. The facial skin is black. Males throughout the distribution, but particularly in Suriname and northern Brazil, show “age related” color variations (transitions), where the pelage of juveniles to subadults is similar to that of females: dark brownish to coppery dorsally with grey-brown-cream stippling on hands, feet, and backs (color depending on the location), bright orange chests (which vary in intensity per region), and shaggy, white, fringy, facial hair that becomes dense, short, and thick in full grown males (Fig. 5). Females. Varying throughout the distribution, fully mature females are brownish to grayish with some cream or offwhite stippling in the dorsal hair, light to bright orange ventral chest hair, and black hands and feet. Females have distinct white to buffy-orange muzzle lines. The variation in females of this species throughout the range is striking and worth further investigation.

Measurements. Since the neotype is mounted with the skull, the following two tables are generalizations of the species based on available data. Table 6 gives average measurements of adult P. pithecia males and females in the museum collections. Table 7 gives examples of cranial measurements for adult male and female P. pithecia.

Diagnosis. This saki is one of the most familiar of all sakis as it survives well in captivity, even outside of South America (ISIS 2012, < www.isis.org>). Its success in temperate climates may be due to its more northerly distribution. They are distinct from P. chrysocephala, in that the latter has bright orange to dark ochraceous facial disks, and obvious, stiff white to buffy hairs along the lips. Further differentiation is seen in the transitional males of P. chrysocephala, which not only look like the chrysocephala females, but are very orange in their faces even when young, as compared to the faces of transitional male pithecia which are white (see “Discussion”).

Distribution. Map 1. Throughout the northern South American countries of Venezuela, Guyana, French Guiana, and Suriname, and in northern Brazil in Roraima, Amapá, and parts of Pará. Those south of this region are P. chrysocephala. This does not reflect precise species delineations (e.g., there are cases apparently referable to P. chrysocephala in Guyana and to P. pithecia south of this initial demarcation), but at present it is not possible to determine if some of these “intermediate” populations are indeed distinct or are color variations of one or the other species (see “Discussion”). In Brazil, it occurs north of the Amazon and primarily east-northeast of the Rio Trombetas, extending north to Roraima, east of the Rio Branco. In French Guiana, Suriname, and Guyana, mostly along tributaries that descend to the coast, some further south into the interior at Wilhelmina Mountains, Keyser Gebergte, and Paloemeu/Tapanahoni rivers in Suriname, and the Rupununi River in Guyana. In Venezuela, as far west as Lago Guri in Bolivar State, south of the Río Orinoco. Their distribution in the Parque Nacional Canaima is unknown but likely as they are in northern Roraima. Their distribution further west and south in Venezuela is unknown.

Specific locations. Appendix I. BRAZIL. Amapá: Rio Jary — Tapuhy, Cachoeira de Sto. Antônio; Mazagão — Rio Maracá, Moreira, Alto Rio Branco; Terezinha — Rio Amupari, Sinturinha, Teomi, Serra do Navio; Rio Tracajatuba; Rio Villa Nova; Rio Caciporé do Ohcana; Municipality of Oiapoque — Villa Velha do Oiapoque. Para: Boiuçú; Bravo — Edo. do Eira; Cabeçeiras do Rio Paru do Oeste — Posto Tirias; Oriximiná — Rio Saracazinho, Cachoeira Porteira; Óbidos — Colonia do Veado; Lago Cuiteña; Patana; Santarém (captive). Roraima: Forte do Rio Branco; São Joaquim. FRENCH GUIANA. Cayenne: Ipousin — Rio Approuague. St. Lamount du Maroni: Mesmond; Saut Macaque sur la Ovaqui; Crique Arouani; Crique Maihonni; Route de Mama a Saint Sabbat. Sinnamary: Saut Tigre; Barrage de Petit Saut; Saisie. GUYANA. Berbice: Essequibo River — Rockstone. Bonasica: Essequibo River — Menarica Creek; Bonnesique Creek. Demerara: coast region; Supinaam River. Mazaruni-Potaro: Barrica; Kaieteur Falls; Kalacoon; Kartabo; Oko Mountains. Upper Takutu—Upper Essequibo: Rupununi River. SURINAME. Brokopondo: Sur River — Gania-Kondre-Bovenkondre; Saramacca River — Loksie Hattie, La Poule. Marowijne: Tapanahoni River — Paloemeu; Moengotapoe. Nickerie: Zuid River — Kayser Gebergte Airstrip; West River — Wilhelmina Mountains; Matapi. Nassau Geb: Marowijne River. Paramaribo: Zanderij. VENEZUELA. Bolivar: El Manaco — 59 Km SE El Dorado, Km 74; La Paragua — Lago Guri (Las Carolinas, Isla Redonda). Orinoco: Orinoco Botanical Garden.

Discussion. There has been a great deal of confusion over P. rufiventer. It was originally included in the Catalogue des Mammifères du Museum National d'Histoire Naturelle, Paris, No. 9 (1803) by É. Geoffroy Saint-Hilaire as Callithrix pithecia, “Le saki ventre roux,” without a number. As of 2007 in Museum National d'Histoire Naturelle, Paris, there is no type specimen available. There is, however, a stand with a label on it in the type specimen vaults which reads: “#457, Saki a ventre roux, P. rufiventer, Guyane, 7-10-1836, male, C.G. 1996–2055.” Perhaps this was the stand for the original type (but if the 1836 date is the date of collection and not that of the making of the mount, then it will not be). In I. Geoffroy Saint-Hilaire's (1851) catalogue, under “S[aki] à ventre roux, P. rufiventer” (1851: 55) only one specimen is listed as having been accessed prior to that date—a specimen collected in Cayenne in 1819 by M. Martin and clearly not the type specimen. Evidently the type has either been lost, or is yet to be identified in the collection. The first use of the name “rufiventer” appears to be by É. Geoffroy Saint-Hilaire in Humboldt and Bonpland's expedition manuscripts (Humboldt et al. 1812; p.357) as part of their expedition to the northern and western regions of South America (described as “Brasils” throughout the text, but they never were that far south; see “Introduction”). Adding to the confusion is the label on the type of P. pogonias in the British Museum of Natural History (No. 42.4.29.7), which is a P. chrysocephala female from Brazil, as: “P. rufiventer, Type, of Pithecia pogonias Gray” (see P. chrysocephala: “Discussion”). By location alone (French Guiana, per É. Geoffroy Saint-Hilaire), I have added those P. rufiventers to P. pithecia. Most of the specimens labeled as P. rufiventer throughout the worldwide museum collections are P. chrysocephala females or subadult males from Brazil, or P. pithecia females from Suriname, southern French Guiana, or Amapá. Since it has not been in use as a valid taxon since 1899, under the provisions of the Code of Nomenclature (1999: Art.23.9.10), I regard P. rufiventer as a nomen oblitum.

There is a possibility of either hybrids or new species throughout the distribution of P. pithecia if the variations in subadult males and females (in Suriname, in particular) prove not to be simple variations within these age-sex categories of P. pithecia and are instead species- or subspecies-defining characteristics. It appears that further north and west to Venezuela, the females are very simple in coloration: grey stippled, shaggy dorsal body hair, orange bellies, and dark grayish-black, mostly uniform faces with white to buffy muzzle lines. In this region, the adult males are generally very black without stippling in the dorsal hairs, with clean white faces. The juvenile and subadult males can be “transitional” in that they resemble females in the grizzling of the dorsal hair and buffy to light orangish bellies, but depending on the region and the age of the individual, the juvenile males can be black with little or no stippling, resembling full adult males.

Moving east and south brings out interesting variations: in areas of Suriname the females are reddish brown and not particularly stippled dorsally; whereas, the subadult males are very stippled with bright orange bellies and white shaggy faces. In contrast to the very white mask of P. pithecia males or the dark orange mask of the P. chrysocephala males, the sakis in Suriname and southern French Guiana appear intermediate, where males often have more orange in the lower halves of their white facial disks as buffy to orangish “cheeks,” or the disk is occasionally entirely light orange in color. Additionally in French Guiana, the animals are smaller and males have distinct lines separating an “outer disk and inner facial disk”, similar to that shown in the original illustration for P. pithecia (Fig. 4). Many females, juveniles, and subadult specimens from Suriname were described originally as distinct species, as P. nocturna and P. ochrocephala, but until further work is done on wild populations to genetically determine if they are distinct in any way or if these specimens are simply off-type coloration from aged curation, they are rolled into P. pithecia.

In Brazil, the females more closely approximate those of P. chrysocephala than those of P. pithecia, and the subadult males are more similar to the Suriname subadult males in coloration. However, in Roraima adult males more closely resemble the P. pithecia type. It is interesting to note that Gray (1870) says P. leucocephala was from Brazil and that “the forehead is yellow when fresh and white when faded by exposure. M. Geoffroy thinks the colour depends on the size of the specimen” (p.59). This is another of the taxonomic confusions early on where they had P. chrysocephala and P. pithecia transitional males standing as types, mixed in with adult males of true species. This reinforces the importance of seeing these animals in the wild. Even with such a well-studied group such as P. pithecia, we have more work to do.

Synonymy

1823 I. Geoffroy Saint-Hilaire. Description Pithecia capillamentosa Spix. Simarium et Vespertiliarum Brasiliensis Species Novae […] Le Voyage dans l–Interieur du Brasil, Monaco, Species Novae p.16, pl. 11. Holotype registered by Kraft (1983) at ZSM. This same mount had many determinations, including P. rufiventer É. Geoffroy Saint-Hilaire, P. leucocephala É. Geoffroy Saint-Hilaire, Simia pithecia and P. pithecia L. (Townsend); ZSM No. 1, juvenile male (fmr. female). Brazil. Nomen dubium (see “Discussion”).

1842 Pithecia pogonias Gray. Ann. Mag. Nat. Hist. Series 1, 10: 256. BMNH No. 42.4.29.7, female. Brazil. Nomen oblitum (see “Discussion”).

1850 Pithecia chrysocephala I. Geoffroy Saint-Hilaire. Comptes rendus hebdomadaires des séances de l'Academie de sciences, Paris 31: 875, Brazil.

1850 Pithecia chrysocephala I. Geoffroy Saint-Hilaire. Description des Mammifères Nouveaux ou Imparfaitement Connus de la Collection du Museum d'Histoire Naturelle, et Remarques sur la Classification et les Caractères des Mammifères, Second Memoire: Singes Americains. Pp.557–559, pl. XXIX of type, Brazil. 1852 Pithecia chrysocephala I. Geoffroy Saint-Hilaire. Catalogue de Primates du Museum d'Histoire Naturelle, Paris, pl. XXXI, p.876, Brazil.

1925 Pithecia monachus lotichiusi Mertens. Senckenbergiana 7(1/2): 17. No. 6692, labeled adult female, fmr. adult male. Holotype, Brazil, Manacapurú. 1925 Pithecia pithecia lotichiusi Mertens. Senckenbergiana 7(3/4): 74. Brazil, Manacapurú.

1939 Pithecia monacha monacha Tate. Bull. Am. Mus. Nat. Hist. 76: 221. Brazil, Amazonas, Jamundá.

1987 Pithecia pithecia chrysocephala Hershkovitz. Am. J. Primatol. 12: 417; in part.

Holotype. P. chrysocephala plate XXIX in I. Geoffroy Saint-Hilaire's original Description des Mammifères Nouveaux for the collection at the Museum d'Histoire Naturelle, Paris (1850) is accepted here and by Hershkovitz (1987) from I. Geoffroy Saint-Hilaire's 1852 publication. I. Geoffroy Saint-Hilaire co-types, mounted skins—one male, one undetermined—reportedly acquired in 1850 by the Museum National d'Histoire Naturelle, Paris, were not there during Hershkovitz's visit (1987), nor were they there as of November 2007.

Neotype. RMNH Leiden (No. 1845(a)): adult male, mount and skull; collected on 15 August 1924 (by unknown), catalogued on 25 May 1930, acquired from the dealers Schlüter & Mass in Halle an der Saale, Germany (C. Smeenk, pers. comm.).

Key specimens. Pithecia p. chrysocephala RMNH Leiden (No. 1845(b)): labeled adult female, likely a subadult male, mount and skull; collected on 27 July 1924 (by unknown), catalogued on 25 May 1930, acquired from the dealers Schlüter & Mass in Halle an der Saale, Germany (C. Smeenk, pers. comm.); Manacapurú, Amazonas, Brazil. Pithecia pogonias holotype, adult female, skin and skull, British Museum of Natural History (No. 42.4.29.7), Brazil. Pithecia (m., p.) lotichiusi holotype at the Senckenberg Museum, Frankfurt (No. 6692), labeled adult female, likely subadult, collected by W. Ehrhardt, 4 July 1924 from Manacapurú “nake Manaos [sic] mittlerer Amazonas, Brazil.”

Type locality. I. Geoffroy Saint-Hilaire stated that the holotype was from “le Brésil, sur les bords du fleuve des Amazones.” Neotype from Manacapurú, Amazonas, Brazil.

Specimens examined. One hundred and twenty-three skins and skulls; photos of living wild and captive animals.

Description. Males. Overall body coloration is black as are hands and feet, similar to P. pithecia, but the facial disk is entirely deep orange or reddish brown. Facial skin is black, bare around eyes, nose, and chin. Upper lip has thick, stiff cream to light orange hairs. Large juvenile to subadult males in some regions are similar to P. pithecia in the same age group where they resemble females (agouti dorsally, including arms, legs and hands, with bright orange chests). Subadult males are more grayish, very mottled compared to females with grey or grey-white agouti, more than just grizzling to make the whole body grayish, but faces are orange to light orange, not fully defined in a disk as they are in adult males. In older juveniles, this facial hair can be quite shaggy to adpressed. The facial pattern can be indistinct and resemble females more closely than males, particularly in younger animals. Females. Dorsal pelage with black to brownish base hairs and some streaking overall, but with tan to buffy arms, legs and tail. The agouti pattern is darker and less extreme than subadult males. Hands and feet are black. The skin around the eyes, nose, and chin is bare and pinkish to grey. Facial hair is in a black horseshoe ring meeting at the top in either a white or orangish to buffy star, and the crown hair over the forehead is whitish or pale and distinct from the dorsal pelage. Orange muzzle lines extend from under the eyes. Their chest and belly is dull to bright orange.

Diagnosis. Pithecia chrysocephala differs from P. pithecia in that males always have bright to dark orange-ochraceous facial disks with contrasting light lips. Females are similar in many ways to P. pithecia females, but tend to be distinct with dark to black facial hair, bright orange malar lines, an orange stripe down the forehead in juveniles and a white star in the center on adults. There are similar females in Guyana P. pithecia populations.

Measurements. See Table 8.

Distribution. Map 2. Only in Brazil, north of the Rio Amazonas, both sides of the Rio Negro, especially in the lower reaches near Manacapurú, east to Faro along the Rio Nhamundá, where populations on the east side appear to be a mix of both chrysocephala and pithecia (see “Discussion”). One specimen in the Berlin collection (BER No. 91313) is clearly an adult male, but is written on both tag and skull to be from the Rio Içá, collected by Lako. If this is true, it is much further west than originally thought. However, this skin was probably collected in the 1920s, and it is unclear if P. chrysocephala still occurs in the region, or if it ever did. The boundaries of P. pithecia and P. chrysocephala are unclear further north in the states of Amapá, Roraima, and Pará, and need further investigation.

Specific locations. Appendix I. BRAZIL. Amazonas: Rio Ica; Rio Negro — Manaus (Km 165, 170, 190), Apuaú - Bocabau Chueiro/Cuieiras, Acajutuba, Porto Mauá, Igarapé do Bolivia, Yavanari, Iaunari, Tabocal; Kastuaria Mirim - Rio Purus(?); Iranduba; Itacoatiara; Itapiranga; Rio Solimões - Manacapurú Rio Itabani nee Atabani; Rio Uruba; Silves; Uatumã - UHE Balbina, Vista Alegre, São Sebastião; Aniba - Igarapé Zinho, Rio Angusta; Lago do Serpa; Lago do Canacary. Pará: Rio Erepecurú - Cachoeira do Tronco; Rio Amazonas - Faro, Rio Piratucu, Serro do Espelho, Nhamundá.

Discussion. The confusion in this species lies with females and juvenile/subadult males. Females in both P. chrysocephala and the Suriname P. pithecia have exaggerated genital swellings with a distended clitoris (see “Introduction: Females”). In preserved skins and mounts this tends to have the appearance of a scrotum and a small penis.

Pithecia capillamentosa. The type of P. capillamentosa Spix in the Zoologische Staatssammlung München, Munich, Germany (ZSM, No. 1) is either a small juvenile male P. chrysocephala from Brazil, or possibly a juvenile male P. pithecia from eastern Brazil, state of Amapá, or even French Guiana as some taxonomists have determined. There are multiple tags on the type in the ZSM: 1) P. capillamentosa Spix, Simia pithecia Lin., P. leucocephala, female or male juv.; 2) P. leucocephala Geoff., Brasilien Spix; 3) P. capillamentosa Spix, Typus, Cayenne, ?sex; 4) P. pithecia L. (Townsend), (?viewed in July) “Saki pithecia (J. Saki Lajuru?—hard to read German script), Catalog J. Akademie, 1816; 5) P. rufiventer Geoff, (hard to read German script, name of journal) Suppl. I, v. 222, Suppl. V, 101, Annal I, Adult female; 6) P. capillamentosa Spix, Simia. Bras., sp. nov. 1823, v. 16, Tab. XI; and 7) a hard to read ZSM tag with what appears to be reference to a (Wagler B- T- hard to read German name), J. Amphib. v. 6, ann. 4, v. 7 with no species det. Pithecia capillamentosa remained a valid taxon and was referenced by several authors Elliot (1912), Cabrera (1957), Hill (1960), and Napier and Napier (1967)), but as it is clearly a juvenile and thus a confusing specimen, given the difficulty of distinguishing juveniles of P. chrysocephala and P. pithecia, the name is best ranked as a nomen dubium (as suggested by C.P. Groves, pers. comm.).

Pithecia pogonias. The type of P. pogonias in the British Museum (Natural History), London (BMNH 42.4.29.7) is an adult female P. chrysocephala resembling those throughout most of the range in Brazil. It is a good specimen, representative of females, with skull, but since the name has not been in use for a valid taxon since 1899, under the provisions of the Code of Nomenclature (1999: Art. 23.9.10), it ranks as a nomen oblitum (as noted by C.P. Groves, pers. comm.). It remains, however, a key specimen as an example of a P. chrysocephala female.

The type of P. lotichiusi in the Naturmuseum Senckenberg, Frankfurt (NMSF 6692), is an adult/subadult female (based on the robustness of the skull), and was named later than P. chrysocephala, and thus ranks as a junior synonym. It is retained here as a key specimen.

The original determination for this species per the holotype set forth by I. Geoffroy was as P. chrysocephala. Hershkovitz (1987) and others considered it to be a subspecies (P. p. chrysocephala) to fit into the two-group scheme. I re-elevate it to species as it is distinctly different from P. pithecia in phenotype, skull morphology, and range. There may be hybrid zones throughout the northern range and into Suriname and the Guianas per discussion in the P. pithecia section.

The Faro sakis appear to be a mix of both chrysocephala males and pithecia; it is interesting that there are also some larger juveniles that appear to have whiter, shaggier faces in the collection from that site with very orange-faced adult males. More work in this region needs to be done to determine if these animals are hybrids, a separate subspecies, different species, or color morphs of chrysocephala.

Synonymy

1823 Pithecia hirsuta Spix, J. B., von. Simiarum et Vespertilionum Brasiliensium Species Novae. F.S. Hübschmann, Munich, pp. 14–15, pl. 9. Pithecia hirsuta, p.14.

Holotype. Adult male mounted specimen with skull intact (No. 19) at the Zoologische Staatssammlung München (ZSM), Munich, Germany, placed by Spix (1823), and as illustrated in the Simiarum. Currently labeled as “syntype” by ZSM.

Syntypes. Mounts at ZSM placed by Spix (1823), Nos. 15, adult male, and 14, juvenile female. Described currently by the ZSM as “syntypes.”

Type locality. The forests between the Rio Negro and the Rio Solimões, Amazonas, Brazil. As for all Spix types a region is reported, not a specific location.

Key specimens. Skins and skulls for adult males, BMNH Nos. 27.3.6.3 and 27.8.11.23 collected by W. Ehrhardt, 29–30 September 1926 (tag note on 27.8.11.23: “Topotype of P. inusta Spix”, which is incorrect (cf. P. inusta types), and adult-subadult females BMNH Nos. 34.6.14.4 and 34.6.14.5. Subadult male from the Rolando Aquino collection (AQ 29.1.82) now in the Museo de Historia Natural, Lima, Peru (UNMSM).

Specimens examined. Sixteen skins and skulls, including photo references of living animals in Brazil and Colombia. Localities of BMNH adult male specimens: Brazil, near the rios Solimões (Tabatinga, on the frontier with Colombia) and Negro, along the Rio Tonantins (possibly near where the original Spix types were collected). BMNH adult females were collected in Brazil, Rio Içá, “Lago do Caraira” (an oxbow lake on the south side of the river, in the upper reaches, as per Paynter and Traylor [1991]; likely it has a different name today). Rolando Aquino collected a subadult male north-east of the Río Napo, along the left side of the Río Tamboryacu, northern Peru (now in the UNMSM).

Description. The most uniform and the plainest of the sakis with very little dichromatism between adult females and males. Males and females are similar in coloring, where the pelage is black with some stippling, but in most cases the whitish stipple is short and not as abundant as in other species. Both have very white hands, and the tail is longer than the head and body. Males. Males with dominantly blackish agouti to brown-agouti head, blackish to brownish chest hairs, and a black chest ruff. The upper section of the chest from the neck to the ruff is obviously bare where the scent gland is positioned. The lips and malar stripes are clearly visible, white to cream and thick. Small bare spots of pinkish to light-colored skin can be seen above the eyes where the rest of the facial skin is predominantly black. Defler (2004) adds, “Males are slightly larger with longer tails, and have a black scent gland or thickened “bulbous sack” under their necks. Males have black testicles and a bright pink penis.” (This accurately describes features of many of the species in Pithecia as the penis is pink, the testes are black, and the scent glands can vary in distinctiveness). Females. Females have more pelage stippling in general than males, but compared to other sakis it is sparse. Females have the white malar stripes as in males, but lack the white hairs across the lips so evident in the males. The hair around the face is blacker than agouti and not as short and tight against the face as in males. The skin on the face is pinkish around the eyes and above the muzzle and the chin, which is black as it is in the males.

Measurements. See Table 9 for Spix's measurements of the holotype, and Table 10 for two adult male key specimens in the British Museum of Natural History (BMNH).

Diagnosis. Pithecia hirsuta is distinct from P. monachus in that it is quite plain, something monachus got the reputation for probably because of É. Geoffroy Saint-Hilaire's simplistic and cryptic original description (cf. P. monachus). Whereas P. monachus adult males have adpressed brown hairs that cover the entire face, and in juvenile males the face can be whitish, P. hirsuta juvenile males have a dense agouti ring with contrasting white malar strips and obvious white lips, nearly resembling the adults. The females of the two species are distinct as well: those of P. hirsuta have very black facial disks and nearly resemble the male hirsuta, only lacking in the white lips, but the female P. monachus has soft, shaggy brown and white facial hair with white malar lines.

Distribution. Map 3. Pithecia hirsuta is distributed in the “wedge” of rivers formed by the Rio Negro to the east in Brazil, north of the Rio Solimões in Brazil and Peru, north of the Río Napo in Peru, and south of the Río Caquetá in Colombia (the Rio Japurá in Brazil). It is not known how far west they occur or where the boundary with P. milleri is, although P. milleri is found in La Paya National Park, Colombia, and probably the Zona Reservada Güeppí in Peru. Pithecia milleri appears in the Cuyabeno-Aguarico region in Ecuador to the exclusion of P. hirsuta (L.K. Marsh, pers. obs.). In Colombia, it occurs in the national natural parks of Amacayacu, Puré, and Cahuinarí for a total of 1,868,380 ha (Defler, 2004 originally referenced as P. m. monachus). In Brazil, it is found south of the Rio Japurá and north of the Rio Solimões-Amazonas, but how far to the Rio Negro in the east they are found is unclear as Spix's type locality is imprecise, suggesting only they are from the swathe between the rios Negro and Solimões.

Specific locations. Appendix I. BRAZIL. Amazonas: Rio Negro/ Solimões — Tabatinga, Rio Tonantins; Rio Içá — Lago do Caroira. COLOMBIA. Amazonas: Río Igara-Paraná — La Raicilla Ravine, La Chorrera; Amacayacu National Natural Park; Río Cahuinari; Bravo Ravine; Río Cotuhe; San Jose del Encanto. Puré National Park: Caño Mateo, Caño Arapa, Caño Esperanza, Quebradón El Ayo; Curare—Los Ingleses Indigenous Reserve: Caño Curare, Caño Agua Blanca, Caño Zumaeta; Camaritagua Indigenous Reserve/ Vereda Madroño: Caño El Boliviano. PERU. Loreto: Río Napo (north) - Río Tamboryacu.

Discussion. The Spix specimens are losing hair, but their corresponding illustrations were drawn when the types were fresh are useful as combined type information. All of the Defler (2004) information used for P. monachus monachus accurately describes P. hirsuta. P. monachus does not occur north of the Amazon River (cf. P. monachus). AMNH 71802, collected by A.M. Olalla on 21 November 1925 is credited with the location of “Boca Río Curaray,” as part of the expedition that was indeed in the area in previous months. It is likely, as it appears to be a P. hirsuta male, that it was collected on the other side of the Rio Napo near that location.

Synonymy

1914 Pithecia milleri. J.A. Allen, New South American Monkeys. Bull. Am. Mus. Nat. Hist. 43: 650.

1987 Pithecia monachus milleri Hershkovitz. Am. J. Primatol. 12: 424–425, in part.

Holotype. Adult male in the American Museum of Natural History, New York, USA, No. 33876 collected by Leo Miller, 8 July 1912 (Collection No. 878), for whom the species was named (Allen, 1914).

Topotype. Juvenile male, No. AMNH 33877, collected and designated by Leo Miller, 9 July 1912, determination by J.A. Allen.

Type locality. Colombia, Department of Caquetá, near Morelia (alt. 700 ft) at the head of the Río Fragua.

Key specimens. Adult male, FMNH No. 70635, collected by P. Hershkovitz, 3 Jan. 1952, determination by Hershkovitz as P. m. milleri, Colombia, Caquetá, Florencia, Montañita, 400 m. Adult female, IAVH No. 0601, collected by H. Chiriví and J. A. Mora, 8 March 1973 from Colombia, Putumayo, Puerto Leguízamo, Caucaya stream, Miguel Velásquez farm, jungle entering by Limoncocha, north-west of Puerto Leguízamo. Note: “Breast with milk, infant capture. Group of five individuals in a swamp area.”

Specimens examined. Thirty-five skins and skulls, wild and captive photo references including those observed and photographed by L. K. Marsh in Cuyabeno, Ecuador.

Description. In overall pelage, males and females are more grizzled than P. hirsuta, and females in particular are noticeably grayer in the face and pelage. Males. As per Allen's (1914) description of the male type: “Upperparts, limbs and tail black, the hairs with long pale yellowish [white] tips; face sparsely clothed with short whitish hairs; front half of head mars brown, the hairs short and course; underparts thinly haired, foreneck naked; hairs dark brown with whitish tips on the throat and belly and with yellowish brown tips over the pectoral region; hands yellowish white, feet whitish grizzled with black.” Subadult and juvenile males have the soft, very brown “mars” coloration of the type as described by Allen, but it is more adpressed in adults making a denser brown band around the face. Males have white hairs along the malars and lips as in hirsuta, but also have white under the eyes and often above as well, which is present in hirsuta but not as obvious. Overall pelage is much more grizzled than in hirsuta, including darker “sleeves” on the forearms where there are patches of brown infused in the black; likewise in the “cuffs” of the hindlimbs. The ruff is not obviously brown or lighter, but can range, as in hirsuta, from a lighter tan to black. Females. Faces are shaggier and whiter in general, sometimes with a distinct white band across the forehead. Longer, shaggier white malar lines, and indistinct white across the lips. Also with whitish hands and feet, less distinct in general than hirsuta.

Diagnosis. Males and females are dichromatic and more distinct from one another than P. hirsuta males and females. Pithecia milleri is distinct from P. hirsuta with males much more grizzled throughout the pelage, and much browner faces and forearms than P. hirsuta. Pithecia milleri females are grayer overall than P. hirsuta females, but particularly gray in the face contrasting with the very black P. hirsuta female faces.

Measurements. See Table 11, including original measurements from Allen (1914).

Distribution. Map 4. In Colombia, Pithecia milleri is distributed from the foothills up to 500–700 m surrounding Florencia, east to the area of La Macarena, and south of the Rio Caquetá to at least Puerto Leguízamo. It is unclear where the boundary with P. hirsuta lies beyond Mecayá. According to Defler (2004), it occurs in La Paya National Natural Park (422,000 ha), Colombia. South of the Río Putumayo it extends into Ecuador north of the Río Napo, but it is unclear how far it occurs east between the Napo and Putumayo. It is possible since Miller's sakis have been recorded in Cuyabeno, Ecuador, that they could be found across the border west into Peru in the Zona Reservada Güeppí, north of the Napo. To date, there have been no records of them in that region.

Specific locations. Appendix I. COLOMBIA. Caquetá: Río Peneya; Río Caquetá - El Infierno - Puerto Santander; La Murelia; Florencia - Montañita; La Tagua - Tres Troncos. Meta: La Macarena National Natural Park. Putumayo: Caucaya Stream - Puerto Leguízamo - Miguel Velasquez Farm/Limoncocha; San Antonio - Mocoa; Puerto Limón - Indigenous Reservation El Picudo - Villa Garzón; Río Mecaya; Caquetá River - El Hacha Stream (between Putumayo and Caquetá). ECUADOR. Sucumbios: Río Aguarico - Lagarto Cocha; Shushufindi; Cuyabeno - Laguna Garza, Laguna Grande, lodges, Río Cuyabeno.

Discussion. Pithecia milleri was described as a distinct species separate from P. monachus by Allen, but Defler (2004) and Hershkovitz (1987) treated it as a subspecies of P. monachus. It is here elevated back to a species. An Olalla specimen, AMNH (No. 71816) collected 25 January 1926, was misplaced by Hershkovitz (1987) as being a female from Peru at “Boca Lagarto Cocha”, and perhaps he was the one who changed the tag on the specimen from “Ecuador” to “Peru.” While it is not entirely incorrect to place it in Peru as the location is near the border with that country, with further investigation, Wiley (2010) discloses: “The Río Lagarto Cocha is a small tributary of the Río Aguarico, itself a tributary of the Napo. The current international boundary between Ecuador and Peru follows the lower Aguarico and the Lagarto Cocha. […] Carlos Olalla's letter stated that it took three days to canoe up the Río Aguarico to reach the Lagarto Cocha. […] Carlos had nothing to say about where his party camped or collected at the mouth of the river. Lagarto Cocha, despite its name (“cocha” means lake in Quechua), is a small black-water river, less than 50 m wide that meanders southward through hilly terrain. Recent satellite photos show the mouth of the river at 0°39′18″S, 75°15′4l″W, but it appears that this mouth is recent, formerly having been about 2 km SW″ (Wiley, 2010, p. 37). While males north-east of the Río Orteguaza appear to be P. milleri, the females, at least in museum collections, almost resemble P. hirsuta with blacker faces than the more typical P. milleri of Ecuador and southwestern Caquetá. As with many saki species, it is possible that the colorations change according to age. Thus, more research is needed on these populations to determine the boundaries with P. hirsuta.

Synonymy

1812 Simia monacha É. Geoffroy Saint-Hilaire. Rec. Obs. Zool. Anat. Comp. (Humboldt expedition). Vol. 2, p.359.

1813 Pithecia monachus. É. Geoffroy Saint-Hilaire. Ann. Mus. Hist. Nat. Paris 19: 116.

Holotype. Skull-in mount MNHN No. 447, Type 92, 2005–928, juvenile male. Other information on mount: “Cabinet de Lisbonne, Brasil 1808. É. Geoffroy St.-Hilaire 1812, Ann. Mus. Hist. Nat. Paris 1. XIX, p. 116.”

Key specimens. Males: AMNH No. 11132, collection no. 210, adult male, skin and skull, collected by D. W. Fleck, 1995–1996 from Peru, lower Río Galvez, eastern side, at Nuevo San Juan. FMNH No. 87002, adult male, skin and skull, C. Kalinowski, January 1957 from Peru, south of the Río Amazonas, Río Maniti, east side, at Santa Cecilia. MPEG No. 1828, adult male, skin and skull, skin and skull, J. Hidasi, November 1960 from Brazil, Rio Javarí, Estirão do Ecuador. Females: MNHN No. 448, catalogue no. 1664, subadult female, skull-in mount, collected on the Castelnau et Deville Expedition, 1867 (which could be the mount date, not the expedition date as it was much earlier) from Peru, Río Yavarí, no specific site. FMNH No. 88861, adult female, skin and skull, collected by C. Kalinowski, September 1957, from Peru, Río Yavarí-Mirim, no specific location. AQ 04.6.82, juvenile female, skin only, collected by R. Aquino, 4 June 1982 from Peru, upper Río Orosa on the west side near the confluence of Río Chontay. BMNH No. 27.3.6.4, adult female, skin and skull, collected by W. Ehrhardt, 10 September 1926 from Brazil, upper Solimões, at Santa Rita, likely present-day Santa Rita do Weil.

Type locality. Geoffroy Saint-Hilaire's (1813) type description says, “Habite…le Brésil?” (p. 117). Others have placed it with Spix's Pithecia hirsuta north of the Rio Solimões, but the type specimen itself has an indeterminate locality.

Specimens examined. Eighty-one specimens, skins, skulls, and photos of living animals from Peru: ríos Yavarí-Mirim, Lago Preto, Tamashiyacu-Tahuayo by M. Bowler, Quebrada Blanco, Río Tahuayo by E. W. Heymann, ríos Tapiche and Tahuayo by various tourists; in Colombia, Leticia area by Juan Manuel Renjifo Rey and Glenn Perrigo, and captives or pets by R. Aquino, L. K. Marsh, and tourists in area of Iquitos.

Description of holotype. Because of the vast confusion about this specimen in particular, I here give detailed information about it for future investigators. Geoffroy Saint-Hilaire (1813) wrote: “Pelage varié par grandes taches de brun et de doré: poils bruns en grande partie et des l'origine, et roux-dorés vers l'estremité: chevelure rayonnante de l'occiput et aboutissant au vertex” (p. 116). He did not provide measurements of this animal, nor an illustration in his publication of it. His description can be applied to several species of Pithecia, both male and female, but mysteriously does not describe the type that was presumably placed by him. The original text from Kühl (1820) repeats in Latin basically what Geoffroy Saint-Hilaire described, but it too in no way describes the specimen that is there now. It does, however, read like it might describe the subadult “transitional” males (see Description, below): “Pilis longissimis, densissimis, ab apice inde bruneis, apicibus tantum extimis ochraceis; ad capitis latera autem et sinciput adpressis, paucioribus, pallide cinerascente ochraceir; in fronte media longitudinaliter haud divisis, occipitis radiantibus, confertissimis. Manuum pilis brevibus, adpressis. Species omnium minima. In Museo Parisiensi” (p. 45). And Gray in 1870 describes monachus in the catalogue for the BMNH as black, grey-washed; hairs very long, harsh, white-tipped; head and crown of male yellow, of female white” (p. 59), which is significantly closer (but see Discussion, below).

If the type specimen is used as the definitive reference, then the one presently in the Paris vaults is quite different from all original references. Gray (1860) refers to the P. monachus type as “from a very young specimen in bad shape” (p. 230) and in 1870 Gray repeats: “The specimen described by Geoffroy was young and in very bad shape,” which sadly does correctly reflect the current type. However, J.A. Allen (1914) vaguely described P. monachus as having a “forehead and crown white,” which approximately agrees with the type as it appears today.

Thus, the problem with the Paris type is many-fold: it is a small juvenile, it was in poor condition as far back as 1860, over time it faded from sunlight hitting the front of the specimen possibly changing the defining coloration (previous curation had the types in clear glass cases for public viewing without regard to natural light or protection from fading, C.P. Groves, pers. comm.), and it could be a female. Hershkovitz (1987) cited the type as “MNHN Paris, No. 447, sex indeterminable, mounted specimen, skull in skin” (p. 423). If the type is in fact from Brazil, there are juveniles with a white band around the face within the monachus range. The confusion remains as far as the type is concerned as there are both young juvenile males and females in that region with white bands around their faces. As an example, MPEG 30768 from Atalaia do Norte, Rio Javarí, Brazil, is labeled “male”, but it is a juvenile female (Figure 33). It is clear from this study that the holotype has value for historical purposes only, not for taxonomic reference.

Description. There is a great deal of variation in the color of the juveniles, both male and female, and the adults. It appears, based on skull morphology, that these sakis in particular live a long time, and their coloration changes over time. In particular, older adult males tend to have some of the most pronounced sagittal crests of all the saki species.

Males. Adult male dorsal pelage is black with very little stippling overall. The forearms and across the chest tend to have longer whitish tips, with the chest ruff having dark roots, but light tan to orangish tips. Juvenile and subadult males are more grizzled in general, with the ruff more obvious and stippling across the arms at chest height make the ruff seem more extensive. The hair from the whorl on the nape of the neck does not extend as far forward over the face as in some sakis, and is darker with less stippling, especially in subadult males. Both adult and juvenile males have black wrists. The hands and feet are mottled black and white, growing whiter with age, but the hairs are subterminally black. The adult males' faces are very brown, and are darker the older they get. The hair is adpressed especially in older animals, with a definite crease up the forehead ending in a star or “pit” between and just above the eyes. In the skulls of older animals, there is a pronounced sagittal crest. Malar lines are white and vary in width depending on age as the white below the cheek line can blend to the malars. There are fine white hairs along the upper lip as well. The facial skin of the muzzle is black with fine whitish hairs sometimes present. In subadult males, the face is white with only a band of brown up the centerline of the forehead. As these animals age, they are similar to adult female P. inusta, and become “two-tone” where the top band of their faces is brown and the bottom white. In some males, the white diminishes to just wide malar lines, and in others it remains white at the lower part of the face. Juvenile males, depending on age, have a white head-band with some brown forming down the center. In very young males, the brown can be nearly across the forehead with the rest of the face white. One of the striking features in older males is the multifolded glandular throat sack, which is more pronounced in this species than any other saki. Females. The overall pelage of females is similar to that of males in that it is black with some stippling throughout. Females in general tend to be slightly more grizzled, especially in the forearms. The hair of the nape whorl extends forward further than in males, but gradually so, it is not as noticeably abrupt as in males. Young, breeding age females can have very loose brownish to whitish mottled faces. Older females have soft, loose dark brown foreheads with shaggy white below, looking “two-toned”. Malar lines are shaggy and white and in older females can blend in with the white lower cheek hair. The ruff on females is darker brown or black, with occasionally lighter tips, that are less extensive than in males. They have less extensive throat glands. In older females, hands and feet are whiter than those of males.

Measurements. None for the type specimen. Table 12 for key specimens of the species.

Diagnosis. Unlike inusta, older adult male facial hair appears more as a band around the face, with some hair filling in toward the muzzle, rather than the face finely covered entirely in hair. Also distinct is the glandular throat sack, and clearly bifurcated forehead muscles on either side of a notable sagittal crest seen in the skulls of some individuals. Faces are much darker brown overall in adult males in P. monachus than P. inusta, and adult female monachus tend toward the brown-topped forehead with the white below, as opposed to the nearly entirely white of adult female inusta.

Distribution. Map 5. In general found in the interfluvial areas between the rios Solimões, lower to middle Ucayali, and lower Yavarí in Peru extending south to at least Sarayacu/Serra do Divisor, and the lower reaches of the rios Javarí to Juruá in Brazil. In Peru: Río Tahuayo region including the Reserva Comunal Amishiyacu-Tahuayo and Quebrada Blanco, north to Río Maniti and Río Orosa, Río YavaríMirim and Río Yavari region, including, Quebrada Esperanza, Río Galvez, north-east to San Fernando, and Leticia/Tabatinga. Tourist photos and specimens of sakis in the ríos Tahuayo and Tapiche are possibly of subadult males or even of P. inusta males. If these animals prove different, it may be because of introgression from P. inusta in the Río Ucayali watershed coming up from the south (cf. “Discussion” below). In Brazil: Benjamin Constant, Santa Rita do Weil, Atalaia do Norte, and Estirão do Equador.

Specific locations. Appendix I. BRAZIL. Amazonas: Rio Javari — Estirão do Equador, Atalaia do Norte; Benjamin Constant — Mata Esperança do Município, Rio Quixito — Seringal, Boa Vista (Gondino), José Veiga; Santa Rita; Tabatinga. PERU. Loreto: Río Yavari; Río Yavari Mirim — Mariscal Ramon, San Fernando, Quebrada Esperanza, Lago Preto; Río Maniti — Maynas, Santa Cecilia; Pebas; Río Amazonas — Orosa, Río Chontay; Río Galvez - Nuevo San Juan; Río Tahuayo — Quebrada Blanca, Tahuayo Lodge, Tamashiyacu; Río Tapiche — Puerto Punga, San Salvador; Iquitos.

Discussion. The “Monachus Mess” has been snowballing for almost 200 years. I read all available early references regarding the description of monachus, including É. Geoffroy (1812, 1813), Kühl (1820), Spix (1823), Gray (1860, 1870), Elliot (1913), J.A. Allen (1914), Lönnberg (1938), and Tate (1939). All of them were confused and conflicting about what Geoffroy meant. Spix (1823) simply ignored monachus altogether, clearly naming and referencing hirsuta and inusta, animals that can still be identified today (cf. P. hirsuta and. inusta). Gray (1870) was not only displeased with the Paris type, but he was also confused about what monachus was by rolling it into synonymy, among others: I. Geoffroy's description of monachus from the Castelnau Expedition, Spix's P. hirsuta, Poeppig's P. guapa and Lesson's two P. nocturna (both of which are P. pithecia transitional juvenile males), and his own P. irrorata saying, “the [specimen] in the British Museum, figured in the ‘Voyage of the Sulphur,’ has the face quite bald. This is now shown to be accidental, as the others, more lately received, have white hairs on the face” (p. 59). We know now that he was right about irrorata, that there actually are bare-faced animals, it is not that the hair has been rubbed off (cf. P. irrorata).

Lönnberg understood the problems best, and tried to give an historic rationale: “P. monachus was named in 1812 by É. Geoffroy St. Hilaire (Ann. Mus. Nat. Paris XIX, p. 116). Although the general characteristics of this monkey are indicated, it has not been sufficiently clear which of the existing races or subspecies is to be considered as the typical one. In consequence of this the specific name quoted has sometimes been used in a rather collective sense, and several new names have also been published. Spix introduced two new names 1823 (Sim et Vesp Bras), the types of which were fully described and also pictured on plates, so that they with full certainty may be recognized vz. hirsuta (i.e., p. 14, PI. IX) and inusta (i.e., p. 15, PI. X). Both these subspecies are represented in the present collection, and as it is not proved, whether any of these can be regarded as identical with the typical monachus, the names of Spix are used.”

Like Spix, Lönnberg never attaches any specimens to monachus, instead he accurately supports Spix's clearly defined species (and adding a new one of his own, cf. P. napensis) leaving us to believe that while Geoffroy indeed was looking at a saki monkey, which saki monkey has been bantered about for generations. Complicating this issue is the fact we have no record of who actually selected the Paris type, nor where it was actually collected. It is exactly as Lönnberg described; that monachus, up to this publication, has been used in a “collective sense,” a dumping ground, of unknown or unnamed sakis. And based on Hershkovitz's last saki work, he fell prey to the same condition. Hershkovitz (1987) stated, “Consistent size and cranial or dental differences between species of the P. monachus group have not been found” (p. 410). This problem for him likely arose from the fact that he lumped several different species and sex/age classes into not only the “monachus group,” but into the monachus species itself (cf. Table 2 “Introduction”).

The clearest, earliest example of what adult P. monachus looks like is Plate 3 in Castelnau et al. (1855) depicting an adult male holding an infant with a portrait of a ‘female’ although the female here is so vague it appears to be more similar to a transitional subadult male or a P. inusta female (Figure 23). The animals depicted in it are from Peru. It is of note that Etienne Geoffroy Saint-Hilaire's son, Isidore, is credited with the species determination in this image and was a member of the expedition into Peru and Brazil. It is this image that allowed P. monachus to remain as a recognizable species, and is what I used to determine adult key specimens.

One of the distinctive characteristics in the P. monachus adult males is their conspicuous, multi-folded glandular throat sack (Figure 28b). It is unclear whether the throat sacks on both males and females are used for scent marking for territory or sexual signaling, although as compared to other New World primates, sakis are not particularly strict territory defenders (pers. obs.). Pithecia sexual behavior has been studied (Setz and Gaspar, 1997; Lehman et al., 2001; DiFiore et al., 2007; Norconk, 2006; Thompson and Norconk, 2011; Thompson et al., 2011), but there are no studies on the size of the glandular areas or excretion rate volume of male glands in relation to sexual advantage, if there is one. Furthermore, while it is clear that at least some sakis signal mates using lingual gestures (tongue moving rapidly in and out of the mouth; Figure 30) similar to other New World primates (for example, Alouatta, pers. obs. for A. pigra, Jones [ ] for A. palliata), it is unclear whether subadults of P. monachus (or any sakis for that matter) initiate sexual partners more frequently with visual cues, such as lingual gestures, as compared to older males with extensive throat glands who may predominantly use scent to attract mates. The function of both visual and olfactory cues in saki sexual behavior needs further investigation in all species.

The deep confusion over P. monachus was not just over its appearance, but its location, as Geoffroy confusingly gives us “Le Brésil?” without any further details, and, as has been mentioned, other authors (including Hershkovitz) simply presumed that he meant what Spix later reported for P. hirsuta. Hershkovitz (1987) was another in a long line who placed monachus thus: “Amazonian region in Ecuador, in Peru the basins of the ríos Huallaga, Ucayali, and Purus in the departments of Amazonas, eastern Huanuco, Pasco, Loreto, and Ucayali, then east into Brazil to the west (left) bank of the Rio Juruá in western Amazonia and Acre; altitudinal range between 50–1500 m above sea level” (p. 423). And while he at least got the right species within those boundaries, other authors presumed this location information to mean that monachus was also in Colombia (Defler 2004) and worse, throughout Ecuador, Peru, Bolivia, and Brazil, all the way to the Rio Tapajós (BDGEOPRIM 2002). It is no wonder we have such divergence in genetic results, and in any kind of comparative anatomy for projects involving “P. monachus.”

With more extensive analysis and considerably more data, it may be shown that P. inusta and 7? monachus are the same species with a huge variation in pelage coloration and skull morphology. Likewise, it may be shown that there are even more species in the Ucayali/Yavarí/Juruá region. The sakis in the lowermost Ucayali, in particular in the ríos Tahuayo and Tapiche, are included in monachus for now, although it is unclear if these animals, photographed by tourists in the region, are subadult monachus males or are adult inusta males. If inusta occurs primarily in the Ucayali drainage and monachus in the Yavarí/Javarí, there could be a “mixing point” somewhere in the north as well as somewhere around Sarayacu/Serra do Divisor in the south. Regardless, the designations of P. inusta and P. monachus as distinct in this publication represent an understanding that there are differences in the sakis of that region, that we need to be careful about creating a new “dumping ground” of species such as monachus was historically, and that “something is going on” in these populations that is worthy of further research.

Synonymy

1824 Pithecia inusta Spix, J. B. von. Simiarum et Vespertilionum Brasiliensium Species Novae. 1823. F. S. Hübschmann, Munich, p. 15–16, pl. 10.

1987 Pithecia monachus monachus Hershkovitz, P. Am. J. Primatol. 12: 422–424, in part.

Holotype. Plate only (Figure 1), original mounted specimen has been lost or destroyed (R. Kraft, ZSM, pers. comm. 2007).

Type locality. Spix (1823) does not specify where the mounted type was collected, not even by country, in his original publication (cf. Discussion below).

Key specimens. MNHN No. 449, subadult male, skull in mount, is similar to the original holotype, this one collected by Castelnau and Deville in 1867 as part of their South American expedition. UNMSM No. 29, skin only, collected by Koepcke in 1949, Peru, near Huanuco, Río Llullapichis, Estación Biológica Panguana. BMNH No. 28.5.2.42, adult male, skin and skull, 18 September 1927, from Peru, upper Río Ucayali, Chicosa, and No. 28.5.2.43, subadult female, skin and skull, 3 August 1927, from Peru, upper Río Ucayali, Cumeria. Both collected at 1500 feet above sea level (457.2 m), and on the Godman-Thomas Expedition (Thomas (1928: 253). And AMNH No. 239853, adult male, skin and skull, collected 28 June 1963 by M. D. Tuttle from Peru, Pasco, Oxapampa Province, Nevati, Mission, 900 feet above sea level (274.3 m).

Specimens examined. Forty-nine specimens, skins and skulls, photos of living sakis by R.A. Mittermeier, J. Vermeer, L.K. Marsh, and tourist photos in the region.

Description. The classic tight, adpressed white faces seen in older museum mounts are confusing at best, as some may be young males and others adult females, the two being very similar. Thus, the following description takes into account the full range of variability among not only adults of this species, but of the females and juvenile age classes. Both males and females have a “diamond” in the center of their foreheads just above the eyes, but in the adult female it tends to be more distinct and less haired than it is in the males.

Males. Dorsal pelage is black with light stippling across the back in juveniles to subadults, more overall in older animals, where the grizzling is short and light initially and increases in length and volume in older males. The ruff is darker brown at the base with light tan tips in younger animals, but can become brighter with buffy-tannish or nearly orange tips in older males. The chest is otherwise sparsely covered in black hair. The forearms in males are sparsely covered in a pattern that for some looks like a triangle up the arm, where half or more is not stippled at all. Wrists tend to be black or without grizzling. The hands and feet are white, and hind feet can have a distinct dark ‘V’ on the older animals.

In adults, the face is entirely covered in short, closely pressed tan to off-white hairs with white, short-haired muzzle lines that are more or less distinct as they blend with the rest of the facial hair. The muzzle is bare, with dark skin, and there are scattered short white hairs along the lips. The transitional males have a very tightly pressed white face and this look can last into subadult ages such that quite large animals appear to have solid white faces like older adult females. All older males in general appear to have a line up the forehead likely caused from muscles beneath the skin forming an indented “line”. Subadults in the northern reaches of the range (Río Tahuayo/Rio Tapiche) have whitish faces with a tan line down the center of the forehead, over the “indent” (cf. P. monachus Discussion); whereas, individuals in the south appear to be more uniform in their burnished color as they get older. There is evidence in some skins that this tan line “spreads out” over the face over time making it the tan-white-burnt color of adults.

Lönnberg (1938) agreed with the original Spix's (1823) description of P. inusta saying: “The crown and the forehead, the cheeks from below the eyes and the sides of the head are densely covered with short, very stiff erect hairs which are directed somewhat forwards, or so on the sides more or less downwards, but are not adpressed. The colour of these hairs is somewhat variable, in some the crown is more pale brownish and others more greyish, and in both cases the color fades downwards and on the sides to more or less whitish [...] Mustache always whitish, short white hairs on lips” (p.8).

Females. The dorsal coloring in adult females is very grizzled, often with a light tan colored ruff, and a mostly white to tannish face, similar to males. Forearms tend to be densely covered with white grizzling and white wrist cuffs (lacking in the males). Faces of adult females are similar to males, although they are longer overall and whiter in general. They also do not tend to have the strong muscular indentations up the forehead as in males. Juvenile females have many transitional “looks” from shaggy, loose hair covering the face that is greyish to brownish to turning shaggy white and eventually a closer white/tan that covers the face similarly as in males. The whorl of hair on the nape of the neck extends longer and in more “bangs” in the females than in males, where it appears to be mostly drawn back and not coming as far forward over the face.

Measurements. See Tables 13–15. A mounted, skull-in specimen, UNMSM No. 54E is an interesting example of an adult female (based on the genitalia of the mount), which looks very similar to Paris 449, a mounted, skull-in subadult male. Canine length on UNMSM 54E is: upper left = 7.8 mm and upper right = 7.9 mm.

Diagnosis. The species most similar to P. inusta is P. monachus, which likely shares part of the northern limits of the range, particularly between the ríos Amazonas/Yavarí in the north-eastern corner of Peru. Pithecia inusta is clearly distinct from monachus and all other sakis in the ontogeny of the juveniles, both females and males. Juvenile male inusta have closely pressed white faces that cover all of the face, and juvenile females have gray-black, sometimes brownish, faces that become white, whereas juvenile male and female monachus tend to be two-toned brown and white, with adult female P. monachus often retaining that coloration.

Distribution. Map 6. In Peru, mostly in the Río Ucayali watershed, where collections appear to have been from both sides of the river from Sarayacu south, especially in the upper reaches where it narrows toward the Río Urubamba/ Río Tambor split. They are in the foothills along the Río Pachitea below Oxapampa, particularly east of the Pachitea along the Río Llullapichis, Monte Alegre, Puerto Victoria, and Nevati/Mission. See “Discussion” below for sakis in the upper Rio Juruá region in Brazil and the upper Río Pachitea in Peru. Population may reach as far north as the ríos Tahuayo and Tapiche in the lower Ucayali watershed (cf. Discussion, below).

Specific Locations. Appendix I. BRAZIL. Amazonas: Rio Juruá — Igarapé do Gordão, Igarapé Grande, João Pessoa. Acre. Upper Rio Juruá — Porongaba, Porto Saids. PERU. Huanaco: Río Pachitea — Monte Alegre, Tabalosos, Pachitea. Pasco: Oxapampa — Nevati Mission, Puerto Victoria; Río Pachitea — Panguana, Río Llullapichis, Río Santiago. Loreto: Río Ucayali — Sarayacu; Contamana — Cerro Azul; Cumeria; Masisea -Tushemo; Chicosa. Ucayali: Río Ucayali; Río Pisqui; Río Urubamaba; Via Utuginia; Río Inuya; Lagarto.

Discussion. “Inustus” in Latin means “burned,” and is an apt description of the adult males with their burnished tan-white faces.

Elliot (1913) was the first to include P. inusta and P. hirsuta as synonyms for P. monachus using the Spix locale of P. hirsuta for all three, even though neither Spix nor Geoffroy give location information for either inusta or monachus: “The type of P. inusta Spix is in the Munich Museum and can in no way be separated from P. monacha. It is full grown in good condition and came from the forests of the Tonantins affluent of the SolimÕes River near Tabatinga” (p. 290) (see “The Monachus Mess,” in P. monachus). And now that there are more data, both from living wild and captive animals and more museum specimens, it is clear the original Spix type is of a large juvenile to subadult male, not an adult. Lönnberg (1938) kept the incorrect location placement, but allowed that P. inusta was indeed a true species based on Spix's type and description. Hershkovitz (1987) and others placed it incorrectly with the P. hirsuta location as described by Elliot (1913). Likewise, there is no specific location indicated for the Castelnau mounted MNHN No. 449 other than Peru, although from his expedition notes it is clear he was never north of the Río Amazonas (Castelnau et al., 1855).

Specimens from the Rio Juruá, Eirunepé (Igarapé do Gordão and Igarapé Grande) in Brazil are included here as part of P. inusta (as per Lönnberg 1938), although further investigation needs to be conducted in this region. In the northern reaches of the Rio Juruá is the much browner, distinctly colored P. monachus. But in the southern reaches of the upper river are these provisional inusta with the older adult males exhibiting heavy skulls, flares on their zygomatic arches, and large sagittal crests, which are closer to monachus than inusta skulls except for the heaviness. The older upper Juruá adult males tend to have faces that are intermediate between P. inusta and P. monachus. The juvenile females tend to be browner than P. inusta juveniles of roughly the same age class, but the adult females are most similar to P. inusta females, not P. monachus. The animals in this region may prove to be subspecies of P. inusta or P. monachus, a distinct species, or color morphs of either.

Sakis in the Río Pachitea region, particularly in the upper reaches (Puerto Victoria and south to Nevati/Mission) are different from the type, where adult males have a softer “medium brown” (intermediate between monachus and inusta) face with white below, and females have blacker faces in general. These are possibly different species, sub-species or color variations, but as there are also animals in the region that are very similar to the type (for example, AMNH 239853), thus they are provisionally included in inusta.

An additional specimen worth mentioning is also provisionally included in P. inusta: SEN No. 16605, skull only, with the compelling label information of “Pithecia monacha Geoffr. M hirsuta Spix, F inusta Spix, location W. Brasilien. Get. 1826 München Mus. gegen Rüppell's Dubl.” Is this the skull belonging to Spix's type in ZSM? Is it one of the Rio Juruá sakis? And it is interesting especially with Rüppell's name associated with it as he was apparently ill and laying low in Africa in 1826, returning to Europe in 1827 ( http://en.wikipedia.org/wiki/Eduard_R%C3%BCppell). It could also easily be monachus or something else entirely. The skull appears to be subadult and likely female.

The sakis in the lowermost Ucayali, in particular in the area of the ríos Tahuayo and Tapiche, are included in monachus for now, although it is unclear if these animals, photographed by tourists in the region, are subadult monachus males or are adult inusta males. If inusta occurs primarily in the Ucayali drainage and monachus in the Yavarí/Javarí, there could be a “mixing point” somewhere in the north as well as somewhere around Sarayacu/Serra do Divisor in the south. Regardless, the designations of P. inusta and P. monachus as distinct in this publication represent an understanding that there are differences in the sakis of that region, that we need to be careful about creating a new “dumping ground” of species such as monachus was historically, and that “something is going on” in these populations that is worthy of further research.

Synonymy

1987 Pithecia irrorata Hershkovitz, P. Am. J. Primatol. 12: 42426, in part.

Etymology. The name was selected for a respected, world-renowned Brazilian primatologist, Dr. José de Sousa e Silva-Júnior (“Cazuza”), who has worked many long years at the Museu Paraense Emilio Goeldi in Belém, now as the Curator of Mammals, where he continues to contribute significantly to South American mammalogy and taxonomy.

Holotype. Subadult male, Museu Paraense Emilio Goeldi No. 37127, skin and skull. Collected by J. Urbechi in várzea, 30 June 2004. Prepared by Arlindo Jr., J. Muniz, and R. R. Silva.

Paratypes. Adult female MNRJ 21055 skin and skull, 28 July 1948, no collector information, but was part of a yellow fever study, “MES Serviço de Estudos e Pesquisas sobre a Febre Amarela.” Large juvenile female MNRJ 3312, skin only, collected by C. Lako, 20 Septl927.

Type locality. Holotype. Uarini, “margem esquerda médio rio Solimões, comunidade Barroso — Reserva de Desenvolvimento Sustentável Mamirauá.” Paratypes. Fonte Boa, Rio Solimões, Amazonas.

Specimens examined. Three (holotype and paratypes), and a photo of a living captive adult male taken in Brazil by  www.tatamazonstock.com and wild adult male by Filipe Ennis.

Description. Both males and females are very plain in their overall pelage with very short, light to absent white grizzling on the tips of their dorsal hairs.

Males. Dorsal hairs black with just the tips dotted in white. Slightly more grizzling over the shoulders and arms, with the white being slightly longer and more extensive. The chest is bare with sparse black hairs, including a minimal black ruff under the bare throat patch. Belly appears to be without pigmentation in the prepared specimen (sub-adult). The arms have brown cuffs at the wrists extending in a brownish ‘V’ down the backs of the hands. On the ankles, there is also a brown cuff ring on top with a less extensive brown ‘V’ down the feet. The hands and feet are off white, largely because they are grizzled with brown/black hairs. The skin on the face of the adult male is black; in the subadult it is black over the nose and muzzle under the chin with a black diamond in the center, otherwise the skin is unpigmented or possibly pinkish. White hairs form a loose arch over the forehead and down the sides of the face. It does not appear adpressed (based on the live photo), but nonetheless is still sparsely haired and the skin (black or unpigmented) shows through it. In both subadults and adults, there is a distinct line through the centerline of the forehead, in adults it is black skin and in subadults it is unpigmented or pinkish. White malar lines are likewise sparse and thin, but the hair continues up under the eyes. The hair on the lips is white, stiff, and quite thick.

Females. The dorsal pelage is similar to that of the males in that it is black with only sparse white grizzling throughout and only on the very tips of the hairs. Older females may be slightly more grizzled, in particular on the forearms and upper shoulders. The hands and feet are white with only very slightly brownish cuffs on the wrists, without a ‘V’ onto the backs of the hands, but with a slight ‘V’ on the feet. The undersides are barely covered in fine black hairs, with a short dark brown ruff with tannish tips. The hair on the head is in a white arch, and while slightly denser and longer than in males, it is still sparse enough to see the facial skin beneath it. The white band dips a bit above the eyes making a diamond in the forehead showing off the bare skin in the middle. The arch peters out below the ears where it is black and white along the sides of the face. The facial skin is black, except for the area just above the eyes where either eyelids or the skin is unpigmented or pinkish. Malar lines are thick, white, and obvious. Lips are lined in white hairs like the males, but are not as dense. Sub-adult/juvenile females are slightly transitional in that they are more grizzled dorsally, have a lighter tan/brown ruff, have a whiter facial arch that covers most of the face to the chin, obvious malar lines as in adults, but pinker or less pigmented facial skin overall.

Diagnosis. These distinct sakis differ from P. pithecia in that the males are not silky black, they are coarsely black with very light tips of white throughout, a short, dark black ruff, and faces with such a diffuse white ring that they are easily distinguished not only from P. pithecia, but from all others with white facial hairs, especially P. aequatorialis which is very grizzled and with a bright orange ruff. They share thick lip hairs with P. hirsuta and P. chrysocephala to the north and northeast of the Rio Amazonas, but differ significantly from both of those species in that they lack the agouti faces of P. hirsuta males and the orange/ochre faces of P. chrysocephala. The females of this new species are unlike any others in the region, including P. monachus and P. albicans, where P. monachus females have brown-white shaggy faces and P. albicans have distinct orange pelage and very short white adpressed hairs covering their faces.

Measurements. See Table 16.

Distribution. Map 7. Appendix I. Not well known, more information is needed. Only in Brazil, and so far only in very northern sections south of the Rio Solimões on either side of the Rio Juruá at Fonte Boa and Uarini.

Discussion. It is never an easy decision to add a new species based on so few specimens. However, upon discovering what is evidently a living representative, I was compelled to call out these unusual animals as distinct.

Synonymy

1987 Pithecia aequatorialis Hershkovitz, P. Am. J. Primatol. 12: 429.

Holotype. FMNH No. 86992, adult male, skin and skull, collected by C. Kalinowski, 3 October 1956.

Topotype. FMNH No. 86994, adult female, skin and skull, C. Kalinowski,

9 October 1956.

Paratypes. Three males: adult FMNH No. 86991, adult FMNH No. 86993, and subadult FMNH No. 86996, skins and skulls, all collected by C. Kalinowski in October 1956.

Type locality. Peru, Loreto District, Río Nanay, Santa Luisa, 160 m; for holotypes and paratypes.

Specimens examined. Twenty skins and skulls; captive photo references from M. Bowler and R.A. Mittermeier; wild photo references from R. Aquino, F. Cornejo, and tourist photos online; and observations in the wild by L.K. Marsh (Monkey Island and Pilpintuwasi, Iquitos, Peru).

Description. Males and females are dichromatic (cf. details in Hershkovitz, 1987).

Males. Have a black pelage overall with long grizzled white tips to the hairs, and chest hair bright orange to ochraceous extending below the armpits and along the sides in a ruff. The hands and feet are white. The faces of the males are black with a dense white horseshoe band of hair around face. The band is adpressed in the holotype, but appears much thicker on living animals. There are two often distinct, white patches above each eye that connect to the white band around the face. They have white malar lines around the muzzle and white lip hairs. Juvenile and infant males have very distinctly white heads with the adult coloration apparent even when very young.

Females. Overall pelage grizzled grayish over black producing an overall grey look. They have an orange ruff that can be as distinct as males, even in juveniles, but not as extensive. Forearms have short brown hairs amongst the black and white. Hair around the face is in a loose grayish white band, shaggier and less formed than in males, with distinct white muzzle lines. The skin of the nose and muzzle is black. Hands and feet are white.

Diagnosis. Pithecia aequatorialis males are distinct from P. napensis and P. isabela sp. nov. in the retention of a dense, fully white arch around the face and a very bright, extensive orange ruff on the chest in addition to the grizzled general body pelage. In P. napensis, the white of the face is only as eyespots above the eyes and as a “headband” between the ears over the crown. In P. isabela sp. nov. the white is focused primarily on the eyespots alone. While both napensis and isabela have varying degrees of orange ruffs, in neither is it quite as extensive as that of P. aequatorialis. Females of aequatorialis are distinct from females of both napensis and isabela in that they are far more white in the ring about the face and much greyer overall than either of the other two species. Juvenile males of all three are quite distinct from each other, with P. aequatorialis being the most obvious at a young age (Figure 51).

Measurements. See Table 17.

Distribution. Map 8. Pithecia aequatorialis is found only in Peru, south of the Río Napo and south (left side) of the Río Curaray to the R´o Tigre in the west (right bank). The northernmost border with P. napensis in this region is unknown. It occurs on both sides of the Río Amazonas at Iquitos, but those on the east side (right bank) are primarily rescues, pets, or escapees at or near Monkey Island Tourist Site or Pilpintuwasi Rescue Center. Additional living photo references are needed to confirm the location in Curaca Corriente in the Reserva Nacional Allpahuayo-Mishana, as the only reference photo to date is of a pelt made into a hat.

Specific Locations. Appendix I. PERU. Loreto: Puerto Indiana; Iquitos; Maynas — Río Nanay — Santa Luisa; north of Rio Tacshacurary — boca Río Machete; Río Napo — Río Tutapisco; Curaca Stream; Reservado Allpahuayo Mishana.

Discussion. All of the P. aequatorialis specimens referenced by Hershkovitz (1987) in Ecuador are P. napensis, except for AMNH No. 98468 from Bassler that Hershkovitz incorrectly assigned to the Río Coca, a northern tributary of the Río Napo in Ecuador. The original label says northern Peru and with further investigation this proved correct. Bassler's expedition was in northern Peru, January–December 1924, and this specimen was obtained between “Guamathra and [“S. Q.”] Tutapischea”, now Tutapisco. This information was obtained from the original expedition records transcribed by Schwartzer at AMNH (unpubl. data).

Aquino et al. (2009) studied P. aequatorialis in north-eastern Peru in the Río Itaya basin (Yanayacu, Seis Unidos, Nauta, Blanquillo), Río Tigre basin (Patria Nueva, Posayo, Coconilla), and the Río Curaray basin (Quebrada Arabela, Sector Curaray). Their observations extend the range of P. aequatorialis in Peru further south-west to between the ríos Tigre and Corrientes, and correctly exclude the area to the north between the ríos Curaray and Napo. I do not dispute their findings; however, they did not take any photographs or samples of the animals throughout their study making it difficult to corroborate this definitively—Pithecia isabela sp. nov. occurs west of the Río Tigre and south of the Río Marañón, and Pithecia napensis extends all the way south from Ecuador to north of the Río Marañón, and likely occurs also west of the Río Tigre watershed (cf. P. isabela sp. nov. and P. napensis).

The phenotypes of P. aequatorialis have seemingly changed over time as well. If one examines the specimens collected by Kalinowski in the 1950s (and there are no other specimens that quite resemble these) with animals alive today, the types are more gracile with closely pressed facial hair; whereas, the present day animals appear to be heavier bodied, bulkier like P. napensis, with the white facial ring not closely adpressed as in the original specimens, but dense and thick and up off of the face—much more like the facial rings on P. napensis, only their white coloration is diffuse and only distinct on the crown (cf. P. napensis). At least per this study, it appears that those animals referred to P. napensis cover a very large territory; this of course may prove to be an artifact of needing more data on where to draw the lines between potential subspecies or color morphs, but P. napensis and P. aequatorialis probably have a hybrid zone somewhere that likely includes the region north of Iquitos/Puerto Indiana. Perhaps this is where we now get the bulkier, thicker white faces of P. aequatorialis today, or the more gracile, adpressed-faced animals of Kalinowski are distinct and will prove to be a separate population if they still exist.

Synonymy

1938 P. monachus napensis Lönnberg, E. Arkiv f. Zoologi 30A (18): 9.

1987 P. monachus monachus Hershkovitz, P. Am. J. Primatol. 12: 423–424. In part.

Holotype. Royal Natural History Museum, Stockholm (RNHM), No. A60-1921, subadult male, skin and skull, collected 13 January 1921 by L. Soderstrom.

Type locality. According to Lönnberg (1938), the type was collected in “Ecuador at Napo river at an altitude of 2000 feet [810 m a.s.l.]” which concurs with Hershkovitz's (1987) information (p. 424). However, the current label reads, “near Napo River and Curaray rivers, west below Macas, approx. 2000 ft, eastern Ecuador” which does not make sense as Macas is in the eastern foothills and the ríos Napo and Curaray are in the north-west. Male key specimen AMNH No. 36468, however, is reported from Baeza, Ecuador, at 3000 feet (914 m), which is not far from the upper reaches of the Río Napo (Allen, 1916), and perhaps is more like the original type.

Key specimens. Adult male, AMNH No. 36468, skin only, collected by W. B. Richardson 1912–1913, purchased from a native collector. Adult male, FMNH No. 41500, skin and skull, collected on 11 February 1922 by R. Olalla from Ecuador, Pastaza, Río Bobanaza, near Montalvo (250 m). BMNH No. 80.5.6.4 adult male, skin and skull, and BMNH No. 80.5.6.5 adult female, skin and skull. Other members of the same group and/or others from the same locale in Ecuador, Río Copataza, 80 miles south-east of Sarayacu: BMNH No. 80.5.6.6 small juvenile male; BMNH No. 80.5.6.7 small juvenile female; BMNH No. 80.5.6.8, large infant male; and BMNH No. 80.5.6.9 small infant male. All BMNH specimens were collected by C. Buckley between December 1877 and February 1878.

Specimens examined. Fifty-eight specimens, including skins and skulls, photographs of living animals from Ecuador and Peru, and observed in the wild in Ecuador by L. K. Marsh.

Description. Males. The most striking feature in the males is that the crown of the head is densely covered with short white hair. The facial disk of dense hair forms a partial disk of white that extends no further than the ears. The disk “fades” into white and gray-black stippling at the bottom. The males have white eyebrow patches leading into the crown that may be more or less distinct and typically blend into the solid white crown on older adults. The skin of the face is unpigmented except for the black nose and muzzle, and appears it could be pinkish in some specimens, but is darker in living animals. Male P. napensis are large and “bulky” with white stippled pelage. They have an enlarged scent gland under the chin that is ringed on the upper chest to the armpits with noticeably bright orange to rusty hair. This coloration of the ruff is conspicuous even at a distance (pers. obs). Male dorsal body hair is black and grizzled with white tips. The amount and length of grizzling varies, but in general is longer on the tail than on the back or arms. Hair on the arms and legs is softer and shorter, and is grizzled on the arms, but not the legs. The hands and feet are white to off-white with hands typically whiter than the feet. The ventrum is mostly naked with sparse grayish-black hair that is light and wispy as it is in most saki species. Subadult males can appear at a distance to only have the white eyespots, but the haloing of the white headband between the ears is present.

Females. The dorsum of the females is black and stippled as in males, but appears grayish overall. The hands and feet are whitish to off-white as in males. The ruff on the females is dark brown with light tan to buffy tips. The facial hair is not short and dense forming a ring as it is in males, but is fine and covers the face, growing mostly downward. The amount of white on and around the face varies with the age of the female, but is never as distinctly organized in a ring as it is in P. aequatorialis females. The facial skin is darker in general than in males, with areas of unpigmented, lighter skin. As in the males, there are distinct white malar lines from under the eyes that follow the curve of the muzzle and continue under the chin to form a half circle.

Diagnosis. Hershkovitz assigned this species to P. hirsuta in 1979 and P. monachus in 1987. Here, I elevate it to species rank. It was correctly described by Lönnberg (1938), although, because of the tenor of the times, he assigned it as a subspecies of monachus as P. m. napensis. However, it is nothing like P. monachus where the adult male faces are brown and not white. And while it is distinct from P. aequatorialis—which is not as bulky in the body, has a far more extensive orange ruff, has a clearly fully white, dense horseshoe ring around the face, and whose juvenile males and females are distinctly different than P. napensis—were napensis to be allied with another species, it would be this one. Pithecia napensis is most similar to P. isabela sp. nov., and sometimes the subadult males of P. napensis can look similar to adult males of P. isabela. But as in the differences in the juveniles of P. aequatorialis and napensis, the isabela juvenile males appear distinct from the others as well (cf. P. isabela sp. nov.). See also “Discussion” below.

Measurements. See Table 18.

Distribution. Map 9. In Ecuador, they are found south of the Río Napo from Coca in the west into Yasuní National Park to the east. The distribution along the Napo further east is not known although likely as populations have been sampled north of the Río Curaray as far south as the mouth where it meets the Río Napo. In the west, specimens have been collected in the foothills west of the Río Coca and west of the Río Napo in Baeza at 914 m and Estribaciones at 1500 m, south to the ríos Bobanaza, Copataza, Pastaza, and Macas in the Pastaza and Morona Santiago provinces.

In Peru, specimens are from the north bank of the mouth of the Río Curaray at the Río Napo. One was photographed by R. Aquino near the Ecuador border on the north bank of the Río Curaray and the left bank of the Río Nashiño. If a specimen from Parinari Caño is from Peru near present day Parinari, north of the Río Marañón (SEN No. 5305, collected on the Bluntschili and Peyer Expedition 24 September 1913, locally called “hirapu”), then P. napensis may extend all the way south to that river. And until there is further data to clarify, animals collected north of the Río Marañón and west of the Río Tigre will be classified as P. napensis, while those to the south will be grouped with P. isabela sp. nov. However, there are samples collected along the Marañón labeled with villages just north that may actually be P. isabela (cf. P. isabela sp. nov.). Sakis collected at the mouth of Río Santiago (right bank, along with Puerto Melendez) will be likewise listed as P. napensis until further data are obtained.

Specific locations. ECUADOR. Napo: Baeza; Río Pastaza. Orellana: Estribaciones; Río Napo; Río Curaray; Macas; Río Cononaco; Yarina; Pamiwa Cocha — Estirão do Equador south Villa Yuca; Río Shiripuno; Yasuní National Park; Napo Wildlife Center; Tiputini Biodiversity Station; Río Maceas. Pastaza: Sarayacu; Río Bobanaza; Montalvo; Río Copataza. Morona Santiago. PERU. Loreto: Boca Río Curaray; Puerto Melendez; Río Tigrillo; Río Marañón. Amazonas: Río Santiago; Río Caterpiza — Villa Huambisa.

Discussion. Heymann et al. (2002) report on their survey results from the Río Curaray: “We saw […] Pithecia monachus only north of the Curaray […] and P. aequatorialis only south of the Curaray. The two species of Pithecia differ in the distribution of white hair in adult males. In Pithecia aequatorialis, the forehead and lateral parts of the face are completely white; in P. monachus, white hair is restricted to irregularly formed patches on the forehead, more similar to the individual in Fig. 16 of Hershkovitz (1987), than to the holotype in his Figs. 22 and 23.” (p. 195). This confusion from Hershkovitz is now corrected with this publication (also see “Introduction, Table 2”), where north of the Curaray we have P. napensis, and south P. aequatorialis; P. monachus is far to the east (cf. P. monachus).

There is a mounted specimen in the Museo Amazónico, Quito, Ecuador, that appears intermediate between P. aequatorialis and P. napensis. It was collected by Shur/Achuar Indians, reported to be from Ecuador, southern Morona Santiago Province. This male has a diffuse white facial ring and less defined eyebrow spots, and is large and bulky, more like P. napensis. As there are few similar specimens, it is unclear if this race is a variation of either P. napensis or P. aequatorialis, is a subspecies, true species, or simply a color variation of P. napensis as the males age where the white becomes more extensive (as it is here presumed). More research on animals in this region is needed.

Subadult male P. napensis can appear very similar to adult males of P. isabela sp. nov. It will be particularly important to sort out animals in the northern Río Marañón, the lower Río Tigre (left bank), and west to Río Santiago to determine where P. napensis and P. isabela sp. nov. share a border, if they do, north of the Río Marañón. Animals from the Río Santiago region, including those from the Río Caterpiza (MVZ Nos. 157794, 1557795) appear to be closer to P. isabela sp. nov. than P. napensis, in particular in the females. They may prove to be different species or subspecies, or included in either P. isabela sp. nov. or P. napensis as color variations.

Hershkovitz (1987) included a specimen (AMNH No. 71817) collected by “Olalla y hijos, 19 January 1926, from Ecuador, Boca Lagarto Cocha” (labeled Peru, but it is right on the border so could be either) in P. aequatorialis, but it is a specimen of P. napensis. What is curious about this specimen is exactly where the Olallas may have collected it. Credited with the same location, AMNH No. 71816, is a juvenile male P. milleri, (cf. P. milleri, “Discussion”). North of the Napo and certainly north of the Río Aguarico is P. milleri territory, and no other P. napensis specimens have been cited there (per this study). As the Olallas were coming up from the Boca Curaray collection site around that time (Wiley 2010) and had another P. milleri in tow (AMNH No. 71802), it is possible they collected it en route on the southern side of the Napo, or it is possible there are P. napensis at the confluence of the ríos Napo and Aguarico near the Peru border. More data needs to be gathered in this region.

There is a Bassler Collection specimen: AMNH No. 98472, an adult male P. napensis without location data on the label. However, per his collection records and the number, it is likely to be an animal he collected near the mouth of the Rio Santiago (unpubl. Bassler Collection records, AMNH). But that location is another “collector's catch-all” of species making it unclear where the specimen was actually from.

Synonymy

1987 P. monachus monachus in Hershkovitz, P. Am. J. Primatol. 12: 422–424. In part.

Holotype. FMNH No. 87001, adult male, skin and skull, collected by C. Kalinowski on 25 November 1956.

Paratypes. Subadult male, skin and skull, FMNH No.122796, collected by P. Hershkovitz, 24 November 1980. Adult male, skin and skull, SEN No. 5304, “Uapu”, collected by Bluntschili and Peyer, 27 August 1912. Adult female, skin only, FMNH No. 86999, collected by C. Kalinowski, 24 November 1956. Subadult female, FMNH 122797, collected by P. Hershkovitz, 30 November 1980.

Etymology. The species is named for an aristocratic colonial Peruvian (recte Ecuadorian) woman named Isabel Grameson Godin des Odonais (Isabel Godin). In 1768, after a 20-year separation from her husband Jean Godin, a cartographer on Charles Marie La Condamine's historic expedition to measure the equator, she mounted an expedition to French Guiana to rejoin him. Her Indian porters, personal African slaves, young Quichua handmaidens, two brothers, a seven-year old nephew, and other travelers amounting to a total of 41 people struck out to cross the Amazon starting from the Río Bamba in the Andes. In less than a month, all of the members of her party either abandoned her or died leaving Isabel alone to struggle in the rainforest for 28 days. She was eventually found by Quichua Indians, who nursed her to health and took her downstream to Andoas, where the priests sent her through the mission system via the ríos Marañon and Amazon. She eventually reached her husband in French Guiana, where they had a happy reunion. Isabel's story of courage and strength is an amazing piece of South American history that absolutely needed proper recognition.

Type locality. The holotype is from Peru, Loreto, along the Rio Samiria at Santa Elena (130 m above sea level). The paratypes are all from the region of the Río Samiria with FMNH Nos. 122798 and 122797 in the Reserva Nacional de Pacaya-Samiria (Base Atun and Biological Station “Pithecia”) and FMNH No. 86999 from Santa Elena.

Specimens examined. Ten skins and skulls, photos of live sakis from M. Bowler, and photos by tourists along the Rio Samiria and in the Reserva Nacional de Pacaya-Samiria.

Description of male holotype. Overall the pelage is black with very light, short white stippling. The “hood” that forms in all sakis with the whirl of hair at the base of the neck is black with little or no stipples. The hands and feet are white with a distinct black “V” extending from the wrists and ankles down the backs of the hands/feet. The forearms have scant to light stippling, but more than on the hind limbs, which are nearly without stippling. The tail is lightly stippled in the manner of the dorsum. The facial disk is dense and dark agouti black to dark brown, with scant white throughout. The eyespots are long, more like stripes, and diffuse with a distinct line between them extending to the dark facial disk. There are white malar lines with some white hairs covering the nose. The skin of the face is black. The ruff is short, orange-ochre, and directly under the throat gland, which is not as distinctly colored as it is in other saki species.

General characteristics. Males. The pelts of some specimens appear coppery almost shiny. The dorsal hairs are black with light, diffuse, short stippling on the back and forearms, very little on the hind legs. The ventrum is sparsely covered with black hairs, the short ruff is on the upper chest right under the throat scent gland and is a dark rusty orange, often very dark, and not appearing more than brown. The hands and feet are white to off-white. White malar lines trace either side of the muzzle. The skin of the face is black, in both, dried specimens and living animals. The facial ring is dark black in adult males and agouti brown in young males, with some light white sprinkled throughout in older animals, but the grizzling varies in intensity or is absent and is mostly sparsely restricted to the forehead if it occurs at all. The facial disk as a whole is dark at the base with light grizzling of whitish or grayish throughout, but lacks a distinct band of white as is seen in P. napensis or P. aequatorialis (cf. “Diagnosis”). The facial hair is short, dense, and straight, and is clearly separate from the hair on the head, with white eyebrow spots over each eye that vary from simple “dots” directly above the eyes (as seen in wild animals at present) to longer stripes up to the darker facial band (as in the holotype), to very diffuse, nearly patternless whitish patches above the eyes.

Females. Overall pelage is similar to males in that it is black, coppery almost in some specimens, with short, light grizzling throughout. There is slightly more grizzling in females than males, but distinctly less than in neighboring P. napensis or P. aequatorialis. The ruff is black with dark brown tips, also right under the throat, but females have a less distinct scent gland. The muzzle lines are thick and distinct and the facial skin is black. The facial disk is black and silky in young females, tighter and black-agouti brown in older females. Adults can have white possible in the facial disk, and with tiny white eyebrows and a small white star just above the eyes. The face is less white than in P. aequatorialis and P. napensis females.

Diagnosis. Pithecia isabela is most similar to P. napensis, with which it shares white patches above the eyes, but differs consistently in the amount of white on the forehead—where napensis is clearly dense and white on the crown and grizzled white throughout the facial disk, P. isabela is vastly more diffuse, with spots smaller and closer to the eyes, and little to no white grizzling around the rest of the face. Pithecia napensis is bulkier, larger, and has a brighter and more extensive orange ruff, whereas P. isabela has a shorter, darker brown-orange ruff that is occasionally a brighter shade. The males in particular are far less grizzled overall in P. isabela, and, in some cases, the grizzling appears nearly absent. Pithecia isabela obviously differs from P. aequatorialis to the north of the Río Marañón in that males have dense, fully white half-circles around their faces with extensive orange ruffs on their chests. Juvenile and subadult males are distinctly different from either P. napensis or P. aequatorialis of the same age. In P. isabela, the faces are plain except for the small white eyespots above the eyes and some diffuse white haloing along the crown. There is no obvious ruff color in the young animals and the grizzling across the back is nearly absent.

Measurements. See Table 19.

Distribution. Map 19. The species occurs only in Peru. The paratypes and most other specimens are from the Pacaya-Samiria region. The paratype was collected along the Río Samiria at Santa Elena. Additional specimens were collected similarly along the Río Samiria at the Base Atún and Estación Biológica “Pithecia.” Mark Bowler reported the species along the Río Yanayacu. Tourist photos show them at Quebrada Sapote near the Ucayali, and collections by R. Olalla suggest they are near Sarayacu on the west side.

Specific distribution. Appendix I. PERU. Loreto: Río Samiria — Santa Elena, Biological Station “Pithecia”, Base Atun; Pacaya-Samiria National Park; Río Yanayacu; Río Ucayali — Sarayacu.

Discussion. This species was originally recognized as distinct and suggested as a new species in 2002 at the Tiputini Biodiversity Station, Ecuador (Marsh, 2004), prior to research for taxonomic revision. In the course of time, the discovery of this new species led to a revision of the genus, and later to placing those in Ecuador with P. napensis upon review of the type specimen in Stockholm.

The Pacaya-Samira National Reserve has become a popular destination for tourists, particularly on riverboats out of Iquitos (for example, Dawn of the Amazon:  http://www.dawnontheamazon.com or La Amatista, a boat run by International Expeditions:  http://www.ietravel.com/central-south-america/amazon-river-cruises). As a result, numerous lodges have sprouted up, some with sakis as freeranging pets. There is some photographic evidence per tourist photos in the region, particularly in the upper reaches of the Pacaya-Samiria near the south side of Nauta, that P. aequatorialis and perhaps P. napensis have been brought from the north side of the Río Marañón. It is unclear if these animals ever leave the facilities they were captured for, but if they do, there may be some interesting ramifications for the P. isabela in the area as time goes on.

In some photos by M. Bowler, females seem to be more similar to P. napensis, where overall pelage appears greyish with more stippling than males. Forearms and chest appear tan, mixed in with the greyish-black, and the hands and feet are white. Faces appear more gray, or at least, with more white mixed into the hair surrounding the face. It is possible it is the lighting or the way in which they were photographed that makes them look closer to P. napensis females as compared to the specimens. Bowler's photos were along the Río Yanayacu, very near to the Rio Marañón. It will be important to study animals throughout the Pacaya-Samiria to determine whether or not P. napensis has in fact established on both sides of the Río Marañón, and where the border with P. isabela lies.

Synonymy

1860 Pithecia albicans Gray, J. E. Proc. Zool. Soc. Lond., p.231, pl.81.

1863 Pithecia albicans Bates, H. W. The Naturalist on the River Amazons, vol. 1, Figs. p. 306, 314.

1987 Pithecia albicans Hershkovitz, P. Am. J. Primatol. 12: 431, for further synonymy.

Holotype. British Museum of Natural History, No. 60.4.16.3, adult male, collected by H. W. Bates, 1862.

Paratypes. Hershkovitz (1987) reports “syntypes” of skins and skulls at BMNH of a male, a female, and “young”, but it is unclear which numbers are assigned as none are designated as such currently. Specimens available: Nos. 27.8.11.17 (M), 34.6.16.10 (F), 26.5.5.6 (F), 26.5.5.5 (F), 26.5.5.3 (M), 26.5.5.4 (F), 26.5.5.2 (M), 27.8.11.18 (M), 26.5.5.1 (M), collected by W. Erhardt between 1925 and 1927. All are undisputed representatives of the species, and can be accepted as types per this taxonomy. Additional specimens from Erhardt's expedition are in the Museum fur Naturkunde, Berlin (Nos. 35768, 35769, 35770, 35308, 35767, 46146).

Type localities. Holotype: Rio Tefé, Tefé Lake, south bank of the Rio Solimões, Amazonas, Brazil. Syntypes: Amazonas, Rio Solimões, Rio Purus, Ayapuá.

Specimens examined. Forty-two museum specimens, skins and skulls; wild photo references from A.D. Johns, and photos of captive sakis by R.A. Mittermeier; and observation of a captive specimen by L.K. Marsh (São Paulo Zoo, São Paulo).

Description. These sakis are the most distinct in the genus due to the obvious coloration and larger size. Males and females are very similar, with predominantly orange to blonde pelage, which in adults is lighter dorsally and darker ventrally on the heads, arms, legs. Ventral chest/belly hairs are sparse. The back is black extending from the nape of the neck to the tip of the tail. The tail is entirely black. The wrists often have black short hairs, the hands are off-white, and the feet off-white. Adult males and females have large throat scent-gland patches ringed in a light orange ruff that is more pronounced in males.

Males. The facial skin is mostly black, with small “bare” pinkish skin patches above the eyes, distinct white to cream eyebrows, and stiff white to cream upper lip hairs. The muzzle has dispersed whitish hairs, not in a distinct line. Ventrally, adult males have testicles that are light cream to buffy, often with black speckles, and on their kneecaps, what appear to be calluses (obvious on intact specimens both living and stuffed, Fig. 70).

Females. The facial skin is black, also with white to cream eyebrows but with white malar stripes that often extend out in short white to cream hairs to the cheeks. Older females have more white covering the face than juveniles or subadults, and may include a short ring of white to off-white hairs around the outside of the face, making the face in some cases appear to be nearly entirely covered in white hairs.

Diagnosis. Pithecia albicans is the “most derived” (sensu Hershkovitz [1987], p.431) of all saki species in that they are the most distinct in pelage. They share some of the buffy characteristics of P. vanzolinii, but as Hershkovitz (1987) notes, “the differences in coloration [...] are wide, and the gap between the incompletely known geographic ranges of the two taxa are also wide” (p. 431).

Measurements. See Tables 20 and 21.

Distribution. Map 11. Pithecia albicans occurs only in Brazil. Hershkovitz (1987) places it between the lower Rio Purus and Rio Tefé, south bank tributaries of the upper Rio Solimões-Amazonas, Amazonas, Brazil. Per the specimen localities studied for this revision, this distribution is accepted specifically noting collections near Lago Tefé, Lago Ayapuá, Lago de Boia, and Jaburú on the left bank of the Rio Purus. Johns (1985, 1986) observed P. albicans in terra firma forest on both sides of the Rio Tefé, and principally at the Ponta da Castanha (3°32′S, 64°58′W) on the eastern side of the lake. They also occur along the Rio Bauana, a tributary entering the lake on its eastern shore (A.B. Rylands, pers. comm, in Johns 1986). Peres (1997) observed P. albicans west of the Rio Tefé, at a locality called Vai Quem Quer, in terra firma on the right bank of the lower Río Juruá, 3°19′S, 66°01′W.

Specific locations. Appendix I. BRAZIL. Amazonas: Rio Purus – Lago Ayapuá, Ayapuá town, Jaburú; Rio Tefé; Tefé town; Lago de Boia; Lago Tefé — Ponta da Castanha.

Discussion. Hershkovitz (1987) and others use the term “buffy” to mean white, off-white, cream, and tan. The definition used here and throughout this text for buffy is based on the original tanned hide definition of “yellow brown” or blonde. Thus, the buffy saki is one which is principally blonde in color, a yellowish brown, not white nor cream per se.

It will be interesting to study the relationship with P. irrorata in the northern reaches of its range in Brazil (west of the Rio Purus and west of the Rio Juruá), P. albicans throughout its range, and Cacajao calvus calvus as a possible evolutionary explanation for why P. albicans is so derived. Strictly from an armchair observation, it seems as though C. c. calvus at some point in time may have bred with the Pithecia in the area. Likewise for P. vanzolinii further south, it will be interesting to study whether the “in between” resemblance of P. irrorata and P. albicans has anything to do with a long ago cross-breeding event or if this extremely different color variation was just a fluke of genetics.

In 2008, Pithecia albicans was classified as Vulnerable (VU) on the IUCN Red List of Threatened Species (Viega et al., 2008). In 1993, Peres reported the population of P. albicans in the upper Rio Urucú to be 8.8 individuals/km². In 2005, Haugaasen and Peres reported an average of 2.2 individuals/km² in the lower Purus watershed at Lago Uauaçù with slightly higher results for terra firma (3.3 individuals/ km²) and igapó (4.6 individiuals/km²), but lower for várzea (0.8 individuals/km²). By all accounts, the population is low if not drastically declining. Research is needed for all areas where P. albicans occurs.

Synonymy

1842 Pithecia irrorata. Gray, J. E. Ann. Mag. Nat. Hist., 1st series, 10: 256.

1844 Gray, J. E. “The Yarke.” List of the Specimens of Mammalia in the Collection of the British Museum, p.14, pl. 3.

1976 Pithecia irrorata. Napier, P. H., Catalogue of Primates in the British Museum (Natural History), Part 1: Families Callitrichidae and Cebidae. British Museum (Natural History), London. Holotype No. 101a; p.69.

1987 Pithecia irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425–426, for further synonymy.

2013 Pithecia hirsuta. Van Roosmalen, M. G. M. and Van Roosmalen, T, 2013. On the Origin of Allopatric Primate Species and the Principle of Metachromic Bleaching. E-book, printed privately, p.63.

Holotype. Hershkovitz (1987) used Gray's (1843) description from vol. 1, color plate 3. No precise specimen was identified, but according to Napier (1976), the type is BMNH No. 101a: adult male, skin and skull.

Type locality. “Tropical America” also said to be “Brazils” (Gray, 1843, p.13), but based on evidence from the actual voyage accounts, it is most likely that the type was procured in Peru (see “Discussion” below).

Key specimens. FMNH No. 98040, adult male, skin and skull, collected by C. Kalinowski, 23 August 1961 and FMNH No. 93534 adult male, skin and skull, collected by C. Kalinowski, 20 October 1960; both from Peru, Madre de Dios, Río Manu, Altamira, 350–400 m. NNMN No. 409 mount, Catalog Nos. 2 & 6, without collector information and date (on specimen) from “Perou Septentr” (implying northern Peru, but without more details on the collector it is difficult to estimate location, even though truly north in Peru is unlikely). MZSP No. 19683 adult male skin collected by J. Hidasi, 24 May 1968, from Brazil, Rio Branco, Acre, the latter at “Fazenda Campo Lindo.” MNRJ No. 3317, adult female, skin and skull, collected by C. Lako, June 1931 from Brazil, Lago do Mapiocu, west of the Rio Purus, assigned by gazetteer as Rio Mapixi (Paynter and Traylor 1991). NHRM No. 1153, large juvenile female, skin and skull, collected by A. Olalla, 20 October 1935 from Brazil, Rio Purus, Arumá.

Specimens examined. Forty-seven skins and skulls, and photographs of living animals in Peru by T. Gregory and tourist photos, and in Brazil by J.C. Bicca-Marques and F. Cardoso.

Description. Transitional juveniles are not as divergent as they are in some species.

Males. Overall, the pelage is black as in most sakis, with longer bands of white grizzling throughout (“irrorata” means “sprinkled”) that vary in density with the age of the animal and location. For instance, the P. irrorata in Brazil, particularly older males in Acre, appear to be very grizzled with white. A distinct white crown “headband” or “bangs” is present—white tips of the nape whorl—but it is not dense as in Pithecia rylandsi sp. nov. In subadults, this line seems to trace down the sides toward the shoulders. The forearms can be brownish to white depending on the region they are from. Pithecia irrorata in Peru, particularly those distant from the Brazilian border, appear to be much browner on the forearms than those north of the Río Tahuamanu. The hands and feet are white. The faces are “pink” or unpigmented with a dark muzzle outlined by noticeable, but fine, short-haired, white malar stripes. Otherwise the hair on the face is lacking or sparse overall. In Peru, animals collected in the early 1960s near the Río Manu/Río Madre de Dios confluence (at Altamira, which no longer exists as that name) were most similar to the type in their faces, even though they were more grizzled white than the type. The sakis further west as far as the Río Urubamba seem to have darker faces as adults, but resemble the juvenile form. Pithecia irrorata from Acre, near Xapuri and Rio Branco, are more similar to the type in that they have pink faces with black muzzles, but are more grizzled in their pelage. In general, males can have darker facial skin as they age, but it never approaches the blackness seen in P. rylandsi sp. nov., nor does their pelage become as white as in P. rylandsi sp. nov. All P. irrorata males have a pronounced ruff ranging from bright orange to dull orange/light tan. Juvenile males can be browner, with grizzling throughout the dorsal pelage. Their facial skin is “pink-black,” where they appear dark, but not black, with lighter pink highlighting around the eyes that is not nearly as contrasting as it is in P. mittermeieri sp. nov. or P. rylandsi sp. nov. juveniles (of either males or females). There is some light white hair on the face above the eyes, and a light “headband” or bang-line that is so distinct in adults is just forming in juveniles.

Females. Overall, the pelage is similar to that of males, where the forearms are very brownish, but not extensively buffy as in P. vanzolinii. They are browner across the front of the shoulders and chest, but do not have a defined light ruff. Young females have very black facial hair around the face with some white in the center of the forehead and a small amount above the eyes. In older adults in Peru, the faces can become grayer. Females in Acre can have an overall body grizzling that is much whiter, approaching more of a P. rylandsi sp. nov. look, with faces retaining the very black facial hair (see “Discussion”). The skin of the face on young P. irrorata can be pinkish-black, but becomes blacker with age.

Measurements. See Table 22. Additional information: Adult male collected by R. Aquino, (AQ 21.4.88) on 21 April 1988 at San Lorenzo, Peru, weighed 2.5 kg and had an upper right canine tooth length of 10.1 mm.

Diagnosis. Pithecia irrorata differs significantly from the other bare-faced sakis in the region: P. vanzolinii and P. rylandsi sp. nov. Pithecia irrorata lacks the obvious buffy forearms and hindlimbs of P. vanzolinii and the very black faces, extreme extent of white pelage grizzling, and size of P. rylandsi sp. nov. P. irrorata have obvious orange ruffs, something P. rylandsi sp. nov. lacks, but P. vanzolinii loosely shares this feature in that it has a buffy-orange ruff to go with its buffy forearms. And while P. irrorata males in the wild may appear to have dark faces, they are mostly unpigmented except for their black muzzle. Pithecia irrorata females differ from either aforementioned species in that up through the subadult class they have black hair surrounding their faces similar to P. vanzolinii females, but as they age this ring can become grizzled with white, something that does not appear to happen in vanzolinii females. Pithecia rylandsi sp. nov. adult females have nearly naked, black faces with scant black hairs along the sides of the face, and can have white distinct eyebrows and pinkish to grey eyespots.

Distribution. Map 12. In Brazil, P. irrorata follows the west side of the Rio Purus, south to Acre. Specifically there are specimens from Lago Mapiocu (likely Lago Mapixi) and Arumá in the northern reaches, and Rio Branco and Xapuri in the south. While it is tempting to say they are in the entire region between the rios Juruá and Purus, there are not enough specimens to support this, especially as the southern reaches of upper Juruá are where P. vanzolinii is found, and the north-west pocket between these rivers is where P. albicans can be found around Tefe and, as of 1935, at Jaburü on the Rio Purus. It is unknown where the boundaries lie with P. vanzolinii in the upper Rio Juruá watershed or P. albicans in the lower Rio Purus. It is also unknown if P. irrorata is found currently further east of the upper Rio Madeira in the southern-most reaches of Rondônia (see “Discussion”).

In Peru, they are primarily in the Manu region, west of the Río Manu and south of the upper Río Madre de Dios, west of the Manu confluence. They extend to the lower Río Urubamba and are likely to occur in all areas north-east of the ríos Urubamba/alto Madre de Dios/Manu to the Brazil border where, theoretically (as there are no specimens of either in the area) they would meet with P. vanzolinii. More data is needed on sakis in this region. Likewise, it is unclear how far east in Peru they extend, as P. rylandsi sp. nov. appears to be in the Los Amigos/Madre de Dios Conservation Concessions. Of note is an adult male P. irrorata collected by R. Aquino in San Lorenzo, Peru, near the Bolivian border that is very similar to the type. It is unclear where exactly it was collected and on which side of the Río Tahuamanu, but it is clear that somewhere near there is a population of P. irrorata, even though P. rylandsi sp. nov. is just over the border in the Pando (cf. P. rylandsi sp. nov.).

Their full distribution is unknown, but Cocha Cashu in their 2008 mammal census list P. irrorata (and I believe the sakis in this region are mostly likely P. irrorata) as “rare”:  http://www.duke.edu/-manu/Home/list_of_species/mammals.htm.

Specific locations. Appendix I. PERU. Ucayali: Río Urubamba. Madre de Dios — Altamira; Río Tahuamanu — San Lorenzo. BRAZIL. Amazonas: Rio Purus — Lago do Arumá, Lago do Mapiocu/Mapixí. Acre: Xapuri; Rio Branco — Fazenda Campo Lindo.

Discussion. Hershkovitz (1987) describes the history of the type specimen thus:

“The formal description of P. irrorata appeared in Gray's 1843 report on the mammals collected between 1836 and 1842 during the globe-encircling voyage of H.M.S. Sulphur. Precisely where, when, how or by whom the barefaced saki was acquired is unknown. In any event, Gray's 1842 reference to P. irrorata was evidently in anticipation of his 1843 report [Gray, 1843b].” (p.409). In the species account he reported that the holotype was “acquired, probably in early 1836, on the voyage of H.M.S. Sulphur.” and restricted it “to west (left) bank Rio Tapajós, Parque Nacional da Amazô;nia, Pará, Brazil.” (p.426).

However, animals in this area do not resemble the type as closely as the sakis further south. Hershkovitz found no reference to the holotype during the accounts of the voyage of H.M.S. Sulphur, neither did C. P. Groves nor A.B. Rylands (pers. comm.). Hershkovitz reported that the Sulphur stopped in Rio de Janeiro, but I could not find any explorations by Belcher or the Sulphur into South America other than Colonial Peru where it is likely the saki specimen is from. However, I do agree with Hershkovitz that likely the holotype of P. irrorata was purchased live in a market by one of the crew and was brought on board as a pet, if it ever really was on board the ship at all. It is equally probable that “Brazils” was synonymous to “South America” at the time, and does not necessarily put the type in the actual country of Brazil (C.P. Groves, pers. comm.).

In Hershkovitz's 1987 paper, Pithecia irrorata was not clearly defined (except by implication in his dichotomous key, p. 416), and he went straight to the level of subspecies denotation since he added P. i. vanzolinii to the species, as he did in cases where he recognized polytypic species. Much like P. monachus, any bald-faced saki was placed under irrorata in a “catch all” manner. It is interesting to speculate at this point on the features in the irrorata of Peru, particularly those that have browner forearms and darker faces in the males and the black hairs around the face of females, as these features are reminiscent of P. vanzolinii. It is possible that the animals in this region are color morphs, subspecies, or different species as they deviate from type.

There is much work to be done on P. irrorata throughout its range. The animals in the state of Acre appear to have more grizzling in their overall pelage than those in Peru, especially one female: MPEG No. 8926, skin only, collected by Flavio Pimenta, 29 January 1971. It is unknown if P. irrorata gets whiter with age as does P. rylandsi sp. nov., but it does seem clear it never approaches the extreme pelage change that is shown by P. rylandsi sp. nov.

Comissão Rondon sakis. Adding to the mystery of irrorata in Brazil, particularly in Rondônia and Mato Grosso, are collections from the Comissão Rondon (1909–1914), collected largely throughout the Rio Aripuaná/Rio Roosevelt—Rio Jamari region that are poorly preserved, but are worthy of further scrutiny. For now they are provisionally added into P. irrorata (for general appearance and skull features). It is entirely possible that these animals are a legacy population from 100 years ago that are no longer there—except perhaps as contributing to a new species in the UHE Samuel region as possible hybrids (cf. Rio Jamari sakis, Appendix II), or are poorly preserved P. mittermeieri sp. nov. as their location suggests (cf. P. mittermeieri sp. nov.).

Furthermore, it is interesting to note that there are some legacy specimens collected as part of the Comissão Rondon (in 1912) that are at present rolled into P. irrorata. These are a juvenile female, skin and skull (MNRJ 3322) and adult male, skin and skull (MNRJ 3339) both from São Manuel/Rio Teles Pires. Although not immediately easy to determine based on their skulls, as both are broken, by their skins they could easily be similar to the more recently collected animals along the Jacaréacanga highway. It is entirely possible that all of these specimens belong in P. mittermeieri sp. nov. based on location, and are artifacts of the preservation process. Certainly, more research needs to be done both on the legacy material and the present day populations in the region.

Sakis from Jacaréacanga. Sakis collected in the early 1970s immediately after the construction of the BR 230 Trans-Amazonica highway between Itaituba and Jacaréacanga on the Río Tapajós have some interesting variations—which could be due to them being a different species, color morphs, or subspecies, or they may be merely remnant populations of P. irrorata. Specimens at the Smithsonian collected by F.D. Pinheiro on 17 July 1972 at km 19 appear to be most similar to P. mittermeieri sp. nov. based on a hairy-faced female in the group (SMITH Nos. 461705, 461706), but animals collected on 19 July 1972 presumably from the same location or nearby (km 19), likely by Pinheiro as well, are quite different. MPEG No. 8150 is a large female with an incomplete skull, but it appears to be adult. She looks similar to P. irrorata females or subadult P. mittermeieri sp. nov. with very black fur around the face, white bangs, and not very much grizzling throughout the pelage. MPEG No. 21536 is an adult male, which in pelage looks intermediate between P. mittermeieri sp. nov. and P. irrorata, where there is a moderate amount of white grizzling, the forearms are slightly more densely covered, there is a prominent orange ruff—and it has a completely pink face! On km 212, collections on 29 November 1973, by an unknown collector are nearly identical to the km-19 animals—MPEG 8149 another adult male—this one showing some fine hairs on the face in a pattern somewhat similar to P. mittermeieri sp. nov. and another adult female, MPEG 21537 nearly identical to MPEG 8150. There are two small juvenile females also from km 212 in the Smithsonian Collection, likely collected by whomever collected the others in 1973 (or perhaps 1972)— SMITH Nos. 461918, 461919—both juvenile females, both appear to have furry faces, but could be either young of these “other” sakis or of P. mittermeieri sp. nov.

East of the Rio Tapajós. While is it entirely possible that sakis east of the Rio Tapajós are not related at all to P. irrorata or P. mittermeieri sp. nov., one of the two specimens is similar to the Tapajós females: MZSP No. 3632, subadult female, skin and skull, from Fazenda Marau, Santarém (according to E. Garbe in 1920 this was about four hours away from Santarém, in the foothills). And the other: MPEG No. 8152, a juvenile female, skin only, resembles a juvenile form of the pink-faced irrorata-like indeterminate sakis from the Jacaréacanga region.

A great deal more research is needed to determine not only where the legacy animals properly align, but if there are any remaining “true” irrorata left in the southern regions of Brazilian Amazon.

Synonymy

1987 Pithecia irrorata vanzolinii Hershkovitz, P. Am. J. Primatol. 12: 426–427; in part.

Holotype. Universidade de São Paulo, Museu de Zoologia (MZUSP), No. 5491, adult male, collected by A.M. Ollala, 29 October 1936, skin and skull.

Paratype. At MZUSP, there are 22 specimens designated by Hershkovitz as paratypes. I would recommend as an example an adult female No. 5495, skin and skull.

Type locality. Brazil, Santa Cruz, Rio Eirú, a southern tributary of the Rio Juruá, Amazonas, for both the holotype and all paratypes.

Description. Species determination is based on 36 specimens. Males and females are similar in pelage with black dorsal fur stippled in light white to cream grizzle, with contrasting pale yellowish-buffy arms and legs. A buffy ruff matches the limbs on both males and females, but the male's ruff is more distinct. A detailed description is given in Hershkovitz (1987). The descriptions by Hershkovitz and this paper are based solely on museum specimens.

Males. The face of the males is bare with black skin and they have white to cream malar stripes.

Females. Black skin on the face is fringed in soft black hair with a star or snip of white on the forehead generally present, and white to cream malar stripes that appear generally wider than in males. Juvenile females appear to have a much shaggier appearance in the face. Female body hair in general is not as thick or wavy as in males.

Diagnosis. Originally placed by Hershkovitz (1987) as a subspecies of P. irrorata, it is here elevated to species—there are no other sakis that closely resemble it.

Measurements. See Table 23.

Distribution. Map 13. Only in Brazil. Hershkovitz (1987) places them in south-western Brazil, between the south bank of the Rio Juruá and the south bank of the Rio Tarauacá in the south-west of the states of Amazonas and Acre. There are, however, very few documented locations for these animals, which are principally along the south-eastern side of the Rio Juruá, including: Amazonas — Rio Eiru, Santa Cruz, Santo Antônio, Lago Grande; Acre — Seringal Oriente “prox. Tassmatinga” (likely Taumaturgo), and Cruzeiro do Sul (which like Sarayacu in Peru or Manaus in Brazil is a site for multiple species collections).

Specific locations. Appendix I. BRAZIL. Amazonas: Rio Jurua - Lago Grande; Rio Eiru - Santa Antônio, Santa Cruz. Acre: Rio Juruá — Seringal Oriente, Tassmatinga, Cruzeiro do Sul

Discussion. As of this writing, I have not seen a living specimen of P. vanzolinii in captivity, in the wild, or as a photograph. It is possible this species is quite restricted in range or occurs in regions far from human populations or tourism. Every effort should be made to gather as much information as possible to determine its range and conservation status.

Synonymy

1987 P. i. irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425–426, in part.

2013 P. irrorata. Van Roosmalen, M. G. M. and Van Roosmalen, T., On the Origin of Allopatric Primate Species and the Principle of Metachromic Bleaching. E-book, printed privately, p.63.

Etymology. Named for Russell A. Mittermeier, President of Conservation International and long-time Chairman of the IUCN/SSC Primate Specialist Group. Dr. Mittermeier, who has been studying primates since 1970, has now seen more primate species in the wild than anyone, and is particularly fond of the Pitheciidae. His dissertation research was carried out in Suriname from 1975–1977 and included some of the first field work on sakis. He has been particularly influential in Brazil, where he has been working since 1971 and where his efforts on behalf of Brazil's amazing biodiversity have earned him the Order of the Southern Cross from the Brazilian government and the João Pedro Cardoso Award from the state of São Paulo. Given his longterm devotion to Brazil, it is especially appropriate that this Brazilian endemic be named after him. He has also been a long-time friend and supporter of my work, so I am also personally very pleased to name this new saki species in his honor.

Holotype. BMNH No. 27.8.5.9.1, adult male, skin and skull, collected by W. Hoffmanns, 14 September 1906.

Paratypes. MZSP No. 5549, adult male, skin and skull, collected by A.M. Olalla, 8 May 1936. BMNH No. 20.7.14.6, subadult male, skin and skull and MNRJ No. 3315, adult female, skin and skull, both collected by F. Lima, 22 July 1917. INPA No. 5707, subadult (lactating) female, skin and skull, collected by P.S. Pinheiro, 23 April 2005.

Type localities. All in Brazil: BMNH No. 28.8.5.9.1 from “Humaytha River,” Rio Madeira, by Gazetteer considered Humaitá (Paynter and Traylor, 1991). Paratypes: Brazil, Pará, Arua, Rio Arapiuns—south of the Rio Amazonas and just west of the Rio Tapajós. BMNH No. 20.7.14.6 and MNRJ No. 3315 Rio Tapajós, Vila Braga (south of Itaituba, left bank). INPA No. 5707 from the “Município de Novo Aripuanã, margem direita do Rio Aripuanã, Lago do Cipotuba.”

Specimens examined. Thirty skins and skulls, additional photographs from captive and living animals throughout the area.

Description. One of the most variable of all saki species, especially in the “transitional” juvenile males. Males and females both have tan to orange fur between the legs around their respective genitalia, similar to P. pissinattii sp. nov.

Males. Adult male dorsal pelage has long white bands of grizzling throughout the black, and the forearms are densely covered in shorter white hairs. Wrist cuffs are white underneath and grizzled white on top. The hind legs are black on the front half, similar to P. pissinattii sp. nov. males, but become very black and obvious as the males age. The hands and feet are white. The ruff is bright orange, but varies in density and extent among individuals. The hair extending toward the face is white and can become a wide white band in adult animals. And as in P. rylandsi sp. nov. the “bangs” are short over the front of the face, while the nape hairs—varying in intensity of whiteness depending on the age—form a “hole” over the ears making it appear as though there are black streaks coming from the sides of the head. It is more prominent in older individuals whose hair has become increasingly white. Adult males, depending on age, have varying degrees of scattered white hairs along the sides of the face and the eyebrows, and as light malar lines and lips. The extent of the light white facial hair decreases as the animals get older. The skin of the face in older animals is black, in younger animals it is black, but the overall look of the face varies with age. Juvenile males of this species are some of the most striking and transitional of all sakis. Very young males have black face skin with prominent pink eyespots that can extend under the eyes as well. As they age, fine white hairs form over the eyes extending back to a horseshoe of fine white fringe around the face. In some photos of living P. mittermeieri subadult males appear “gray” faced, where the white can seem obvious and extensive (but short and light) and the eyespots not as intensely pink. Depending on the angle of the photo or the way the specimen was prepared and depending on the age, juveniles can appear greyish, coppery-brownish, very black with a white fringe, or colorful with both white hairs and very pink eyespots.

Females. Overall, the pelage is similar to that of males, but generally much less grizzled, especially when younger. As the females age, they become more grizzled overall, but never approach the whiteness seen in P. rylandsi sp. nov. females. Adult females have a darker ruff than males in general, but it sometimes can be a lighter tan tending toward orangish in some individuals. Very young females resemble young males with pink eyespots over the eyes and black faces, but they always have thick, shaggy malar lines, and a fringe of black hair around the back edge of the face, and hairs forming over the brow. As they age they have extensive facial hair that is white, similar to P. pissinattii sp. nov. females. Some subadult individuals have very black fur around the face, and very little pelage stippling. As they age, this hair can go from dark, to loosely white, to more closely pressed to the face as blackish with white in it. The facial skin is black, with scattered hairs over the muzzle.

Diagnosis. Distinct from all other sakis in that the males are almost “clownish” with bright orange ruffs, grayish lightly haired faces as subadults, with traces of hair over the black faces in older adult males, and contrasting very white and black pelage with dense buffy to white forearms. Much older adults can be quite white in overall pelage, but unlike P. rylandsi sp. nov. males, retain their bright orange ruffs. Females in the upper sections of the range south of the Amazon and between the rios Madeira and Tapajós have very hairy faces as compared to the females of P. rylandsi sp. nov.

Measurements. See Table 24.

Distribution. Map 14. Pithecia mittermeieri is found only in Brazil, south of the Rio Amazonas between the rios Madeira and Tapajós, including the Rio Aripuanã drainage. Current populations appear to be concentrated primarily north of Aripuanã in Mato Grosso, although there is some evidence that historically they may have extended at least along the Madeira as far as the Mamoré and even Rio Guaporé (as per Natterer 1829 in Destacamento do Ribeirão and Rio Pacáas Novos, but cf. P. rylandsi sp. nov.). The collection of a male, juvenile male, and infant in “Nova Brasilia, Polonoroeste,” presumed to be Ji-Paraná as per the gazetteers of Paynter and Traylor (1991), in 1985, demonstrates some populations north of the Rio Jamarí as well. Two photos of wild males taken in southern Rondônia suggest they are still in the region. One was a tourist photo in 2010 at Cacoal Selva Park, 470 km from Porto Velho, and the other in Chupinguaia in 2012 (Figure 85). Along the Tapajós, they extend possibly as far as Jacaréacanga and perhaps further south, depending on the presence of other species (see Discussion below).

Specific location. Appendix I. BRAZIL. Para: Rio Madeira - Borba, Humaitá; Parintins - Villa Bella Imperatriz, Lago Andina; Itaituba; Rio Arapiuns - Ama; Rio Humaitá; Rio Aripuanã; Rio Tapajós; Lago do Cipotuba; Villa Braga; Igarapé Auara; Igarapé Brabo; Igarapé Amorin; Limoal; Limontuba; Santarém - Fazenda Mararu. Mato Grosso: Aripuanã. Rondônia: Nova Brasília.

Discussion. The colloquial name for these sakis has generally been “the Tapajós saki,” and it will likely continue to be so even though the species name has been given as P. mittermeieri. The region between the rios Madeira and Tapajós has changed extensively over the last two hundred years. Railways and telegraph lines were introduced at the turn of the 19 th century (instigated by Marechal Rondon). Because of highways such as the BR 364 in Rondônia completed in 1961, the Trans-Amazonica BR 230 inaugurated in 1972, and the devastating colonization and development program Polonoroeste (Northeastern Brazil Integration Development Program) in the 1980s, the forests have gone from complete coverage in Rondônia and Mato Grosso in the 1950s to approximately half or less today (per satellite images). Previous collection sites are now roads and towns leaving little doubt that the primate populations have been severely disrupted and have potentially led to hybrid zones (cf. Jamari sakis, Appendix II).

Likewise, as discussed for P. irrorata, it is not known if historic populations of P. irrorata were somewhere within the collecting routes of the Comissão Rondon, or how far north of the Rio Guaporé P. rylandsi sp. nov. extended originally, but today's distribution is likely very different for all of these species. So while it is possible that P. mittermeieri extended throughout the region between the rios Madeira and Tapajós to the Guaporé, its exact distribution today is unknown.

Synonymy

1842 Pithecia irrorata. Gray, J. E. Ann. Mag. Nat. Hist., 1st series 10: 256. In part.

1976 Pithecia monachus. Heltne P., Freese, C. and Whitesides, G. A Field Survey of Nonhuman Primates in Bolivia, Report, PAHO, Washington, DC. 40pp. 1981 Pithecia hirsuta. Izawa, K. and Bejarano, G. Kyoto Univ. Overseas Res. Rep. New World Monkeys (2): 3, 5,10.

1981 Pithecia hirsuta. Izawa, K. and Yoneda, M. Kyoto University Overseas Res. Rep. New World Monkeys (2): 16.

1987 Pithecia irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425–426; in part.

2013 Pithecia hirsuta. Van Roosmalen, M. G. M. and Van Roosmalen, T, On the Origin of Allopatric Primate Species and the Principle of Metachromic Bleaching. E-book, printed privately, p.63.

Etymology. Anthony Rylands is a Senior Research Scientist at Conservation International, Deputy Chair of the IUCN/SSC Primate Specialist Group, a member of the Brazilian Academy of Sciences, former professor of Vertebrate Zoology at the Federal University of Minas Gerais, and founding editor for the journal Neotropical Primates. He has mentored hundreds of young primatologists, and as coordinator for Conservation International's Primate Action Fund, has directly impacted primate research and conservation the world over. Dr. Rylands is a tireless academic and a dear friend. For all of these reasons and more, I name this species after him.

Holotype. AMNH No. 247669, subadult male, skin only, collected by K. Izawa, 5 December 1980 (listed as female on the label, but face appears male). As type skin does not have a corresponding skull, I include AMNH No. 248723, subadult male, skull only, collected by S. Anderson, 26 May 1982.

Paratypes. FMNH No. 123967, skin only, subadult female. As paratype skin does not have a corresponding skull, I submit KUPRI No. 5973, skull only, collected by M. Kimursa, 8 November 1978.

Key Specimens. KUPRI Nos. 5974 and 5975, both skulls only, collected by M. Kimursa in Bolivia with No. 5975 collected in Mucden, Bolivia on 5 August 1979.

Type localities. Bolivia, Pando Department: Holotype skin AMNH No. 247669 from the Río Tahuamanu, Mucden; holotype skull AMNH No. 248723 from the Río Nareuda.

Paratypes: FMNH No. 123967 from Bolivia, Pando Department, Porvenir, Río Nareuda. KUPRI No. 5973 from Bolivia.

Specimens examined. Eleven skins, 14 skulls, and 40 photos of living specimens (captive, rescued, wild) from numerous sources.

Information about this species comes in part from living photographs of wild and captive animals in Bolivia, Peru, and Brazil. No museum specimens are available from Peru. Specimens from Brazil are provisional pending further investigation. According to R. B. Wallace (pers. comm.), there are no Pithecia specimens in any of the Bolivian natural history museums.

Description. One of the largest of the sakis. Males and females have bare faces with black pigmentation.

Males. Overall pelage is transitional, as it is in most of the genus. When young, the males are black and moderately to heavily grizzled with white throughout. As they age, they become almost entirely white. Their dorsal coloration can be so intense with white that they take on a “skunky” appearance. Males in Peru appear to have almost no black showing through the heavy white hair tips. The forelimbs can be buffy to white, and the hind limbs can have black on the inner leg, but it may not get as black as P. mittermeieri or the Jamari sakis (cf. Appendix II). The hands and feet are white to off-white. The crown bangs are also white in all ages, and can be quite thick compared to other species, and are shorter than the hair coming from the shoulders, making a “hole” where the ears are. This makes for the appearance of black stripes on the sides of the older males' heads. This kind of pattern also appears in very old P. mittermeieri and sometimes in P. irrorata. Males tend to lack or have only very light malar lines, but do have white hair on the lips. They lack an obvious orange or bright ruff as in P. irrorata; instead they have a black or dark brownish ruff. Males retain light to pinkish eyespots over the eyes. Juvenile males are dark “pink-black,” where the facial skin is not as dark as adults and appears reddish-black, but tend toward black even from very young. Otherwise males have plain, extremely lightly furred or non-furred faces.

Females. Adult female pelage can be as intensely white as males and the hair on the forehead, creating “bangs,” can look like a white shield. Their forearms can be more buffy than those of the males, and this blonde-tan can extend across their chests. They do not have distinct ruff coloration, as it is sometimes a light buff or grayish. The black of their faces is deep with reddish undertones, and lacks hair. Younger females can have obvious white eyebrow hair. Very young females can look cartoonish with pink circles around the eyes, a very black face with some black hairs along the sides, and white forming as eyebrows. All ages of females have distinct, white, shaggy muzzle stripes. Young females have grayish faces, sometimes with a ring of light black hairs peripherally around the face with light grey eyebrow spots and light skin around eyes. Adult females retain the light colored eyebrow spots, and, depending on age, may retain some of the light black fringe, but in most cases, the faces are very bare save the bright muzzle lines.

Diagnosis. Easily distinguishable from all other bald-faced sakis by their large size, very black faces, and extreme white grizzling. P. irrorata males have far less white throughout, bright orange ruffs and light faces with dark muzzles. P. irrorata females have black-fringed faces as juveniles becoming greyer faced overall.

Measurements. See Table 25.

Distribution. Map 15. Found in north-western Bolivia, south-eastern Peru, and possibly in the south of the state of Rondônia and the west of the state of Mato Grosso in Brazil.

Bolivia. Found in the western Pando Department, but with some evidence that they may be in northeastern Beni and northern La Paz, particularly north/northwest of the Río Madre de Dios in those areas. Specific locations with verification include: Mucden; north of the Río Tahuamanu, but east of the Peru border; and Municipio de Bolpebra, Provincia Nicolas Suarez. Additional locations reported for Bolivia by various authors without specimens or photos, summarized by R. B. Wallace (unpubl. data) by municipality include: Bolpebra — Buena Vista, Piaou, Rutina, Los Campos, El Refugio, Estación Tahuamanu, and San Sebastian (reported ‘not common’); Filadelfia — Pingo de Oro (‘rare’) and north of Río Tahuamanu, vicinity of Río Muyumanu; Bella Flor — Santa Rosa near Río Abunã, Bella Flor, Río Manuripi, and between the ríos Acre and Tahuamanu; Cobija — Cobija, Belo Horizonte, Tres Corazones, Puerto Rico, Tres Estrellas, Fortaleza; Porvenir — Porvenir along the Río Nareuda and Cocamita; Nuevo Esperanza — Federico Roman (Compartamentos Caiman, Manoa, and Río Negro); and Ingavi — Centro Dieciocho, north of Barraca San Juan de Nuevo Mundo.

Peru. Mainly north of the Río Madre de Dios, likely throughout the Reserva Territorial Madre de Dios, specifically in the Los Amigos Conservation Concession including Río Los Amigos, the Manu Wildlife Center, and Los Amigos Research Center. It is possible, as sakis are reported for Tambopata, that they are also south of the Río Madre de Dios, but this is unconfirmed. Their border with P. irrorata north of the Madre de Dios to the Brazilian border and further west toward the Parque Nacional Alto Purus is unknown.

Brazil. While it is likely that these animals occur east and north of the Rio Guaporé into the states of Rondônia and Mato Grosso, and possibly as far north as the Rio Anari or west to the Rio Jamari (see Discussion, below), to be conservative in this account, their distribution will be restricted to the south (west) of the Rio Guaporé or just north (east) along its Brazilian shores where passable by arboreal primates.

Specific locations. Appendix I. BOLIVIA. Pando: Nicolas Suarez - Bolpebra; Tahuamanu; Mucden; Rio Nareuda. BRAZIL. Rondônia: Cacoal Selva Park; Rio Guaporé; Destacamento do Ribeiro; Rio Anari; Pedra Branca; Ritiro do Veado Branco - Serra do Norte; Guajarí-Mirim. PERU. Madre de Dios: Río Madre de Dios - Manu Wildlife Center; Río Los Amigos Conservation Concession.

More data are needed to determine the borders and range of these monkeys in relation to those of P. irrorata in Peru and Acre, Brazil, and P. mittermeieri north of this area.

Discussion. These sakis are distinct and unmistakable. More data are needed to determine the boundaries with P. irrorata in Peru and P. mittermeieri in the southern Brazilian Amazon. This is one of the few cases in this taxonomy where there is more information on living animals than reference material.

Material collected by J. Natterer in 1829 from Guajará-Mirim, Rio Pacaás Novos (old name, now very near Guajará-Mirim), and Rio Guaporé are juveniles and difficult to place firmly in P. rylandsi; therefore, as they are thought to have been collected on the north bank of the Rio Guaporé, they will be provisionally placed in P. mittermeieri (cf. P. mittermeieri). A female (MZSP No. 22895) collected in Rondônia, Rio Anari, Pedra Branca by D. F. Stotz that appears to be closer to P. rylandsi than to P. mittermeieri, extends the possibility that they were further north into Brazil at least in 1988. More study is needed for the P. rylandsi population in Brazil.

Except for south-eastern Peru and perhaps parts of Pando in Bolivia, it is likely the population is declining throughout the range. While I do not advocate more collections of these sakis, I do suggest we get as much detailed information from photos or from capture-release animals as possible to determine their relationship to P. irrorata and P. mittermeieri.

Synonymy

1987 Pithecia irrorata irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425–426, in part.

2013 P. hirsuta. Van Roosmalen, M. G. M. and Van Roosmalen, T, 2013. On the Origin of Allopatric Primate Species and the Principle of Metachromic Bleaching. E-book, printed privately, p.63.

Etymology. Named for Dr. Alcides Pissinatti, a Brazilian veterinarian, director and co-founder of the Centro de Primatologia do Rio de Janeiro (CPRJ/FEEMA), and Vice President of the Brazilian Academy of Veterinary Sciences. For nearly two decades Dr. Pissinatti has almost singlehandedly pioneered captive breeding programs for endangered Brazilian primates, including the successful breeding and management of the muriqui (Brachyteles) in captivity. Because of his genuine kindness and intellect, and his devotion to the primates of Brazil including Pithecia, I dedicate this species to him.

Holotype. BMNH No. 26.5.5.7, adult male, skin and skull, collected by W. Ehrhardt, 14 November 1925.

Paratype. BMNH No. 25.12.11.2, adult male, skin and skull, and BMNH 27.8.11.13, adult female, skin and skull, collected from Canabouça by W. Ehrhardt, 14 November 1925. INPA No. 4060, juvenile female, skin and skull, from Autazes, collected by Marc G.M. van Roosmalen.

Type locality. BMNH holotype from “Middle Amazonas, Paraná do Jacaré, Canabouça.” Hershkovitz and others assign this location to Lago do Canabouça, just south of the Rio Solimões and between the rios Purus and Madeira.

Specimens examined. Thirty-five specimens, skins and skulls, and photos of living sakis by R.A. Mittermeier, R. Sampaio, and various tourists.

Description. Male and female faces are mostly pinkish to a darker red, tending toward black in older females. Both have distinct grizzling of white throughout the long black dorsal pelage that varies by age in its amount, but compared to other bald-faced sakis, it can be comparatively sparse in some individuals.

Males. Pelage varies, with younger males very grizzled in general, much more like females, and older males less so. The color bands of grizzling in the dorsal pelage vary as well from whitish to buffy-cream to tan, particularly on the forearms and shoulders, often with whiter cuffs at the wrist. The hind legs are often darker, brownish to blackish, sometimes with dark bands lacking the white grizzling. The ventrum is lightly covered in black hairs with a distinct ruff ranging from bright orange to dull orange/light tan, generally with lighter roots. The hands and feet are white. Males have a white crown band of “bangs,” hair that is distinct, but not thick as in other species, that drapes over the forehead. Faces are mostly bare and are a deep reddish that seems almost a waxy reddish/black, even in well-preserved museum specimens. The hair on the face is lacking to sparse in adults, but with white malar lines that extend up under the eyes. Fine hairs above the eyes and around the back edges of the face sometimes remain even on older animals. Lips and muzzles can have sparse indistinct white hairs. In younger males, the faces can have more fine hair, but not nearly as much as the juvenile males east of the Rio Madeira (P. mittermeieri).

Females. Females often have more grizzling in the dorsal pelage than males—the back is tanner than in males, and especially on the arms and legs. Like males, their back legs can be half black along the front edge over the knee. The ruff is present with dark brown base hairs ending in lighter tan to buffy bands. They often have darker wrist cuff bands above the hands. The faces of adult females are covered in white soft hairs that cover the forehead, cheeks, and eyebrows. Females often have light white hairs especially on the forehead, and in older females, a loose white-grey band around the face, with a skin “diamond” in the center of the forehead just above the nose, between the eyes. The white malar lines are more obvious and often shaggier than in males. The skin above and below the eyes is light pinkish, especially in younger females, with the rest of the bare parts of the face black. In older individuals, the bare skin is blacker overall.

Diagnosis. Populations of P. pissinattii collected in 1935 in the region are very similar to those photographed by tourists on the Rio Juma (at Juma Jungle Lodge) as recently as 2009 (Figure 96, 98a,b). Pithecia pissinattii are most similar to P. mittermeieri on the eastern side of the Rio Madeira, in that adult males with darker faces appearing black can in some photos seem similar to P. mittermeieri, especially when coupled with the brighter orange ruffs. However, juvenile and subadult male P. mittermeieri are very different from those of P. pissinattii. The young P. mittermeieri males have very bright pink eye spots that give way to white facial hair that in older subadults makes them have an overall “grey appearance,” and this does not seem to be the case for P. pissinattii. However, it is possible that P. pissinattii may have legacy populations that were on both sides of the Rio Madeira, lending to features we now see in P. mittermeieri, particularly in females.

Measurements. See Table 26. Additional information: One skull measured, BMNH No. 25.12.11.1, subadult male (mm): braincase length 81.8, braincase width 42.1, zygomatic arch width 55.1, outer span of orbits 42.5, nasal constriction 0.74, orbit width — inner 15.2, outer 16.0, muzzle width 24.5, and mandible width 50.1.

Distribution. Map 16. In Brazil only, they are found south of the Rio Solimões in the northern area between the rios Purus and Madeira, including Autazes, Lago do Canabouça, lagos Sampiao and Miguel (including the area formerly called “Rosarinho”), Rio Juma, and one specimen reported from Itaboca, which appears to be a juvenile male P. pissinattii collected from the right bank (NHRM No. A62-1200). It is not known how far south between the Purus/Madeira these animals occur before they meet with P. irrorata, if they do at all.

Specific locations. Appendix I. BRAZIL. Amazonas: Solimões — Parana do Jacaré, Canabouça, Lago Canabouça; Rio Purus — Lago Jari, Itaboca; Autazes; Rio Juma — Juma Lodge; Rio Madeira — Rosarinho, Lago Miguel, Lago Sampaio; Manaquiri.

Discussion. Pithecia pissinattii sakis may occur east of the Rio Madeira, and gene flow with P. mittermeieri may be impacting the population west of the Rio Madeira. More research is needed, however. As with many saki populations in this monograph, the convention of large watersheds is used as a preliminary hypothesis for the separation of species, particularly in their lower, wider reaches.