Neotropical primates

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Title:
Neotropical primates a newsletter of the Neotropical Section of the IUCNSSC Primate Specialist Group
Abbreviated Title:
Neotrop. primates
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v. : ill. ; 27 cm.
Language:
English
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IUCN/SSC Primate Specialist Group -- Neotropical Section
IUCN/SSC Primate Specialist Group -- Neotropical Section
Conservation International
Center for Applied Biodiversity Science
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Conservation International
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Belo Horizonte Minas Gerais Brazil
Belo Horizonte Minas Gerais Brazil
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Primates -- Periodicals -- Latin America   ( lcsh )
Primates -- Periodicals   ( lcsh )
Wildlife conservation -- Periodicals   ( lcsh )
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review   ( marcgt )
periodical   ( marcgt )
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Brazil

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Also issued online.
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English, Portuguese, and Spanish.
Dates or Sequential Designation:
Vol. 1, no. 1 (Mar. 1993)-
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Issued jointly with Center for Applied Biodiversity Science, <Dec. 2004->
General Note:
Published in Washington, D.C., Dec. 1999-Apr. 2005 , Arlington, VA, Aug. 2005-
General Note:
Latest issue consulted: Vol. 13, no. 1 (Apr. 2005).

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University of Florida
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University of Florida
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All applicable rights reserved by the source institution and holding location.
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oclc - 28561619
lccn - 96648813
issn - 1413-4705
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UF00098814:00066


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NEOTROPICAL PRIMATES rfnnftbnfbrfn bfntnbbft n n bfbb bbffb bbn n nbr bffnnnbnn nfnbnbffnnbnntfbf Neotropical Primates A Journal and Newsletter of the IUCN/SSC Primate Specialist Group Vol. 21(1), July 2014ContentsA Taxonomic Revision of the Saki Monkeys, Pithecia Desmarest, 1804Laura K. Marsh .................................................................................................................................................................1 NEOTROPICAL PRIMATES rfntbb

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Neotropical PrimatesA Journal of the Neotropical Section of the IUCN/SSC Primate Specialist Group Conservation International 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USAISSN 1413-4703 Abbreviation: Neotrop. Primates Editors Erwin Palacios, Conservacin Internacional Colombia, Bogot DC, Colombia Liliana Corts Ortiz, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA Jlio Csar Bicca-Marques, Pontifcia Universidade Catlica do Rio Grande do Sul, Porto Alegre, Brasil Eckhard Heymann, Deutsches Primatenzentrum, Gttingen, Germany Jessica Lynch Alfaro, Institute for Society and Genetics, University of California-Los Angeles, Los Angeles, CA, USA Anita Stone, Department of Biology, Eastern Michigan University, Ypsilanti, MI, USA News and Books Reviews Brenda Solrzano, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Ernesto Rodrguez-Luna, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Founding Editors Anthony B. Rylands, Center for Applied Biodiversity Science Conservation International, Arlington VA, USA Ernesto Rodrguez-Luna, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Editorial Board Bruna Bezerra, University of Louisville, Louisville, KY, USA Hannah M. Buchanan-Smith, University of Stirling, Stirling, Scotland, UK Adelmar F. Coimbra-Filho, Academia Brasileira de Cincias, Rio de Janeiro, Brazil Carolyn M. Crockett, Regional Primate Research Center, University of Washington, Seattle, WA, USA Stephen F. Ferrari, Universidade Federal do Sergipe, Aracaj, Brazil Russell A. Mittermeier, Conservation International, Arlington, VA, USA Marta D. Mudry, Universidad de Buenos Aires, Argentina Anthony Rylands, Conservation International, Arlington, VA, USA Horcio Schneider, Universidade Federal do Par, Campus Universitrio de Bragana, Brazil Karen B. Strier, University of Wisconsin, Madison, WI, USA Maria Emlia Yamamoto, Universidade Federal do Rio Grande do Norte, Natal, Brazil Primate Specialist Group Chairman, Russell A. Mittermeier Deputy Chair, Anthony B. Rylands Vice Chair, Special Section on Great Apes, Liz Williamson Vice Chair, Special Section on Small Apes, Benjamin M. Rawson Regional Vice ChairsNeotropics Mesoamerica, Liliana Corts-Ortiz Andean Countries, Erwin Palacios and Eckhard W. Heymann Brazil and the Guianas, M. Ceclia M. Kierul, Fabiano R. de Melo and Mauricio Talebi Regional Vice Chairs Africa W. Scott McGraw, Janette Wallis and David N.M. Mbora Regional Vice Chairs Madagascar Christoph Schwitzer and Jonah Ratsimbazafy Regional Vice Chairs Asia China, Long Yongcheng Southeast Asia, Jatna Supriatna, Christian Roos, Ramesh Boonratana, and Benjamin M. Rawson South Asia, Sally Walker and Sanjay Molur Layout: Kim Meek, Washington, DC IUCN/SSC Primate Specialist Group logo courtesy of Stephen D. Nash, 2002. Front cover: Adult male Pithecia hirsuta, Capar Biological Station, Colombia. Photo by R.A. Mittermeier. is issue of Neotropical Primates was kindly sponsored by the Margot Marsh Biodiversity Foundation, 432 Walker Road, Great Falls, Virginia 22066, USA, and the Los Angeles Zoo, Director John R. Lewis, 5333 Zoo Drive, Los Angeles, California 90027, USA. Printed in Bogot DC, Colombia on July 25, 2014. NP 20_supplement_COVER_k.indd 2 7/25/14 10:33 PM Scope e journal/newsletter aims to provide a basis for conservation information relating to the primates of the Neotropics. We welcome texts on any aspect of primate conservation, including articles, thesis abstracts, news items, recent events, recent publications, primatological society information and suchlike. Submissions Please send all English and Spanish contributions to: Erwin Palacios, Conservacin Internacional Colombia, Carrera 13 # 71-41 Bogot D.C., Colombia, Tel: (571) 345-2852/54, Fax: (571) 3452852/54, e-mail: , and all Portuguese contributions to: Jlio Csar Bicca-Marques, Departamento de Biodiversidade e Ecologia, Pontifcia Universidade Catlica do Rio Grande do Sul, Av. Ipiranga, 6681 Prdio 12A, Porto Alegre, RS 90619-900, Brasil, Tel: (55) (51) 3320-3545 ext. 4742, Fax: (55) (51) 3320-3612, e-mail: . Contributions Manuscripts may be in English, Spanish or Portuguese, and should be double-spaced and accompanied by the text on CD for PC compatible text-editors (MS-Word, WordPerfect, Excel, and Access), and/or e-mailed to (English, Spanish) or (Portuguese). Hard copies should be supplied for all gures (illustrations and maps) and tables. e full name and address for each author should be included. Please avoid abbreviations and acronyms without the name in full. Authors whose rst language is not English should please have their English manuscripts carefully reviewed by a native English speaker. Articles. Each issue of Neotropical Primates will include up to three full articles, limited to the following topics: Taxonomy, Systematics, Genetics (when relevant for systematics and conservation), Biogeography, Ecology and Conservation. Text for full articles should be typewritten, double-spaced with no less than 12 cpi font (preferably Times New Roman) and 3-cm margins throughout, and should not exceed 25 pages in length (including references). Please include an abstract in the same language as the rest of the text (English, Spanish or Portuguese) and (optional) one in Portuguese or Spanish (if the text is written in English) or English (if the text is written in Spanish or Portuguese). Tables and illustrations should be limited to six, except in cases where they are fundamental for the text (as in species descriptions, for example). Full articles will be sent out for peer-review. For articles that include protein or nucleic acid sequences, authors must deposit data in a publicly available database such as GenBank/EMBL/ DNA Data Bank of Japan, Brookhaven, or Swiss-Prot, and provide an accession number for inclusion in the published paper. Short articles. ese manuscripts are usually reviewed only by the editors. A broader range of topics is encouraged, including such as behavioral research, in the interests of informing on general research activities that contribute to our understanding of platyrrhines. We encourage reports on projects and conservation and research programs (who, what, where, when, why, etc.) and most particularly information on geographical distributions, locality records, and protected areas and the primates that occur in them. Text should be typewritten, doublespaced with no less than 12 cpi (preferably Times New Roman) font and 3-cm margins throughout, and should not exceed 12 pages in length (including references). Figures and maps. Articles may include small black-andwhite photographs, high-quality gures, and high-quality maps. (Resolution: 300 dpi. Column widths: one-column = 8-cm wide; two-columns = 17-cm wide). Please keep these to a minimum. We stress the importance of providing maps that are publishable. Tables. Tables should be double-spaced, using font size 10, and prepared with MS Word. Each table should have a brief title. News items. Please send us information on projects, eld sites, courses, esis or Dissertations recently defended, recent publications, awards, events, activities of Primate Societies, etc. References. Examples of house style may be found throughout this journal. In-text citations should be rst ordered chronologically and then in alphabetical order. For example, (Fritz, 1970; Albert, 1980, 2004; Oates, 1981; Roberts, 2000; Smith, 2000; Albert et al., 2001) In the list of references, the title of the article, name of the journal, and editorial should be written in the same language as they were published. All conjunctions and prepositions (i.e., and, In) should be written in the same language as rest of the manuscript (i.e., y or e, En or Em). is also applies for other text in references (such as PhD thesis, accessed see below). Please refer to these examples when listing references: Journal article Stallings, J. D. and Mittermeier, R. A. 1983. e black-tailed marmoset ( Callithrix argentata melanura) recorded from Paraguay. Am. J. Primatol. 4: 159. Chapter in book Brockelman, W. Y. and Ali, R. 1987. Methods of surveying and sampling forest primate populations. In: Primate Conservation in the Tropical Rain Forest, C. W. Marsh and R. A. Mittermeier (eds.), pp.23. Alan R. Liss, New York. Book Napier, P. H. 1976. Catalogue of Primates in the British Museum (Natural History). Part 1: Families Callitrichidae and Cebidae. British Museum (Natural History), London. esis/Dissertation Wallace, R. B. 1998. e behavioural ecology of black spider monkeys in north-eastern Bolivia. Doctoral thesis, University of Liverpool, Liverpool, UK. Report Muckenhirn, N. A., Mortensen, B. K., Vessey, S., Fraser, C. E. O. and Singh, B. 1975. Report on a primate survey in Guyana. Unpublished report, Pan American Health Organization, Washington, DC. Website UNESCO. 2005. UNESCO Man and the Biosphere Programme. United Nations Educational, Scientic, and Cultural Organisation (UNESCO), Paris. Website: http://www.unesco.org/mab/index.htm. Accessed 25 April 2005. (Acessada em 25 de abril de 2005 and Consultado el 25 de abril de 2005 for articles in Portuguese and Spanish respectively). For references in Portuguese and Spanish: and changes to e and y for articles in Portuguese and Spanish respectively. In changes to Em and En for articles in Portuguese and Spanish respectively. Doctoral thesis changes to Tese de Doutoramento and Tesis de Doctorado for articles in Portuguese and Spanish respectively. MSc esis changes to Dissertao de Mestrado and Tesis de Maestra for articles in Portuguese and Spanish respectively. Unpublished report changes to Relatrio Tcnico and Reporte no publicado for articles in Portuguese and Spanish respectively. Notes to Contributors NP 20_supplement_COVER_k.indd 3 7/25/14 10:33 PM

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1 enLaura K. Marshen1en1enGloenben alen en Coennen serenven atienoen n Iennen stitenuen te, Saennen ta Fe, New Meenxen icenoen USA; lkmarsh@global-conservation.orgAbstractFor more than 200 years, the taxonomy of Pithecia has been oating on the misunderstanding of a few species, in particular P. pithecia and P. monachus. In this revision, historical names and descriptions are addressed and original type material is examined. For every museum specimen, all location, collection, and museum data were recorded, and photographs and measurements of each skin, skull, mount, or uid specimen were taken. e revision is based on work conducted in 36 museums in 28 cities from 17 countries in North America, South America, Europe, and Japan, resulting in the examination of 876 skins (including mounts and uids), 690 skulls, and hundreds of photographs taken by the author and by colleagues in the eld of living captive and wild sakis of all species, and through internet searches. Per this revision, there are 16 species of Pithecia: ve currently recognized, three reinstated, three elevated from subspecies level, and ve newly described. Key Words: Pithecia, saki, taxonomy, revision, new species, P. pithecia, P. mon a chusResumenPor ms de 200 aos, la taxonoma de Pithecia ha estado otando en la confusin de unas pocas especies, en particular P. pithecia y P monachus. En esta revisin, los nombres histricos y descripciones son abordadas y el material tipo original es examinado. Para cada espcimen de museo, todos los datos de localidad, coleccin y museo fueron registrados, y fotografas y medidas de cada piel, crneo, montaje, o especmenes conservados en uido fueron tomadas. La revisin est basada en trabajo llevado a cabo en 36 museos en 28 ciudades de 17 pases en Norte Amrica, Sur Amrica, Europa, y Japn, resul tando en la examinacin de 876 pieles (incluyendo montajes y uidos), 690 crneos, y cientos de fotografas tomadas por la autora y por colegas en el campo, de individuos vivos en cautiverio o estado silvestre de todas las especies de Pithecia, y mediante bsquedas en internet. Con base en esta revisin, hay 16 especies de Pithecia: cinco actualmente conocidas, tres reclasicadas, tres elevadas del nivel de subespecie y cinco nuevas descritas. Palabras Clave: Pithecia, saki, taxonoma, revisin, nueva especie, P pithecia, P monachus A T R S M, PITHECIA es-ESDr, 1804

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2 W h at gets us i n to tr oub le is not what we d o n t kno w its w h at we k now f o r sure t h at ai nt so. Mark Twain Introductione saki monkeys of the genus Pithecia are a poorly studied group of Neotropical primates. Found throughout tropical forests of South America from the Guiana Shield, west to the foothills of the Andes in Ecuador and Peru, south to northern Bolivia, and throughout the Amazon Basin in Brazil as far east as Altamira on the Rio Xingu, these secretive, fast moving, medium-sized monkeys (called ying monkeys in many languages) have eluded researchers for decades. ere have been few long-term eld studies, thus creating a huge knowl edge and conservation gap for these unusual animals. Sakis are the smallest of the pitheciines, but are larger than other non-prehensile-tailed platyrrhines (Aotus, Saimiri, Cal licebus, and the Callitrichinae). Adults weigh in at 1.5.0 kg, with a wide range in total body length (250 mm) and tail length (255 mm) reecting the diversity among the taxa (Buchanan et al., 1980; Mittermeier et al., 1981; L.K. Marsh, unpubl. data). eir tails are often 1:1 with the body length, but in many species tails are longer by >100 mm (based on museum specimen data) for both males and females. Size dimorphism is not striking but, depending on the species, females tend to be smaller in weight and overall size. Sakis are characterized by their long, coarse, ufy hair that easily piloerects when they are approached in the wild, making them look bigger than they actually are. e hair on the tail tends to be longer than on the back, but this varies among species, and the tail hair also pufs up as a threat. Hair on the trunk is always longer than that on the forearms and hind limbs, and underbellies are very lightly covered or bare. Coronal hairs are directed forward as a hood, and, depending on the species, overlap the facial region making them appear to have bangs. Both males and females have distinct throat glands for scent marking, with varying development depending on the species; some males have obvious neck folds and others only a bare skin patch (Brumloop et al., 1994; Setz and Gaspar, 1997). Sakis are frugivore/seed predators, but they eat insects such as army ants, as well as spiders, and other arthropods (Bu chanan et al., 1980; Deer, 2004; pers. obs.). Locomotion consists of quadrupedal walking/running, climbing, and leap ing (Youlatos, 1999; Deer, 2004). ey vocalize in grunts, chirps, whistles, and low calls, but can be exceptionally quiet when sneaking away from a perceived threat such as a eld researcher (pers. obs.). All species tend to prefer mature forest that includes variations of ooded forestvrzea (white-water ooded forest) and igap (black-water ooded forest)palm swamp ( Mauritia swamp for some), and terra rma forest (Rylands, 1987; Rylands and Mittermeier, 2009). ey can be found in disturbed habitats and in fragments with secondary forest, but as seed predators of large forest tree species (notably of the family Lecythida ceae), they tend to be in higher densities in mature forest (Norconk 2003). One possible explanation for their very large geographic distribution as a genus and as individual species is their ability to use a range of forest types, from vrzea to igap to terra rma (Ayres and Clutton-Brock, 1992).Previous Taxonomyere has been a great deal of confusion in Pithecia taxonomy. e most recent revision was by Hershkovitz (1987). Later publications (e.g., Rylands et al., 2000; Groves, 2001; Deer, 2004) reiterate Hershkovitzs work (Table 1). Hershkovitz (1987) dened ve species with three of them polytypic. ey were arranged into two groups as follows: 1) Pithecia pithecia Group Guianan region: P. pithecia pithecia, P. pithecia chrysocephala, and 2) Pithecia monachus Group Amazonian region: P. monachus milleri, P. monachus monachus, P. irrorata irrorata, P. irrorata vanzolinii, P. aequatorialis, P. albicans. Hershkovitz was compelled to force everything in the genus into this construct, thus missing key diferences in what prove to be distinct species, including those that may be diferent enough that they should not be grouped together (e.g., irrorata, aequatorialis, or albicans). An interpretation of the specimens identied in Hershkovitz (1987) is in Table 2, elaborating the diculty and confusion he faced while studying these animals. For Hershkovitz and others, Pithecia was a succession of allopatric species with no overlap or chance for mixing. In reality throughout the range, Pithecia species likely show sympatry with other members of the genus, or are parapatric with hybrid zones between them. e use of groupings and subspecies has been suspended in this publication until detailed studies on genetics, behavior, morphology, cur rent distribution, and evolutionary relationships have been completed with the guidance of this new taxonomy. Only then can comparable studies be performed and subspecies accurately determined (Groves, 2004).en Early taxonomists Early taxonomists (1700s through the turn of the 20th cen tury) had a hard time describing sakis because they were so unusual. One of the common mistakes made in describing them was the use of the term beard, as members of the genus Pithecia do not have beards. Since Pithecia was originally included with Chiropotes (the truly bearded sakis), this mistake was common. More often than not, what the early taxonomists were referring to in Pithecia as a beard was their chest ruf, distinct in several species. It is a shame that Pithecia shares the colloquial name of saki (a Qui chua term for Pithecia) with Chiropotes, as the native term

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3 for Chiropotes is cuxi (coo-shiu). Perhaps if we make a dis tinction in the common names for these two, we will stave of confusion later. Early taxonomists and collectors also had no idea about the behavior of Pithecia in the wild. Gray in 1870 called them gregarious, slow, sad, voracious, and vociferous (p.59). And while it was clear Gray never saw sakis in the wild, Bates (1892) had seen one as a pet: e Parauac is a timid inofensive creature, with a long bear-like coat of harsh speckled-grey hair. e long fur hangs over the head, half concealing the pleasing diminutive face, and clothes also the tail to the tip, which member is well developed, being eighteen inches in length, or longer than the body. [] e Parauac is also a very delicate animal, rarely living many weeks in captivity; but any one who succeeds in keeping it alive for a month or two, gains by it a most afectionate pet. One of the specimens of Pithecia albicans now in the British Museum was then living on the property of a young Frenchman, a neighbour of mine at Ega. It became so tame in the course of a few weeks, that it fol lowed him about the street like a dog. My friend was a tailor, and the little pet used to spend the greater part of the day seated on his shoulder, whilst he was at work on his board. It showed, nevertheless, great dislike to strangers, and was not on good terms with any other member of my friends household than himself. I saw no monkey that showed so strong a personal attachment as this gentle, timid, silent little creature (p. 336). Sclater (1879) probably understood best: We know very little of [Pithecia] habits in the wild state, although it appears that they are normally silent (p.174). It has been my experience with captive sakis of various species that they are indeed quite calm, gentle, and allow themselves to be petted by humans.e Trouble with SakisMany erroneously labeled types have been placed in the museums that house Pithecia specimens. ere are a number of reasons for this, including the historic methods of specimen collection and the lack of eld observations; the age of individuals placed as types and the longevity of sakis in the wild; misunderstanding of females and their anatomy; and the business of transitional males.en Historic collection of specimens Until the latter half of the 20th century, the taxonomist in question might have gone to the country of interest for col lecting, but he likely remained behind any real expedition into the forest in favor of a nearby town or city. He would have a team of local hunters or collection experts culling the animals for him, and more often than not, sakis were among the wild life plunder as opposed to animals of targeted interest. us, many specimens have location designations such as Cay enne, Iquitos, Manaus, Sarayacu, or other similar towns even if the animals are from much further aeld or from the other side of a river. is leads to more than one species being described for a site and does not necessarily account for actual distribution. For instance, the abundance of types in French Guiana and from Cayenne in particular likely represents the number of French explorers at the time more than the diver sity of the region. Worse, early explorers (1700s) would simply label locations as Tropical America, South America or Brasils, which may or may not have actually meant the country of Brazil, as Brasils was a generic term for South America 100 years ago. Even some of South Americas most prolic collectors, the Olalla brothers, may not have been as reliable as we would wish to believe by their specimen tag information. Wiley (2010), in an excellent tracing of the history of the Olalla familys work collecting birds in Peru and Brazil, recounts the then (1965) curator for Field Museum Emmet R. Blakes impression of the men: Blakes impression was that Carlos [Olalla] was suspect, but that Alfonso [A.M. Olalla] was not given to actual skullduggery, although generally he didnt bother to indicate which side of a river his [specimens] came from [...] he simply didnt know any better. Blake went on to say that [Philip] Hershkovitz ... bears me out on this and has had exactly the same trouble with monkeys borrowed from the AMNH that you are having with some of our Olalla birds. In fact, Philip [Hershkovitz] admits that in his younger collecting days he also was less than precise in speci fying localities as related to the banks of rivers (p.5). Traditional taxonomists, including Hershkovitz, almost always made their determinations once the samples were back in the home museums. Out of context and far from the forests where the sakis lived, they gave names to ju venile males and all ages of females as though they were adult males of a new species. It has only been in the last decade that information for determinations has been so readily available to a taxonomist in the form of digital photography, international cross-referencing, easy communication with colleagues around the world, and photos of living animalsin the wild and captivitythroughout the range. In the past, memory, notes, verbal descriptions, the occasional painted plate, or later lm photography (which relied heavily on the development process) were the princi pal ways to determine and compare species. Taxonomists working from their home museums, particu larly 100 years ago, worked almost blind when describ ing saki specimens. If they had not been to Paris, London, or Munich to see the actual specimens placed there by tienne Geofroy St.-Hilaire, John Edward Gray, or Johann Baptist von Spix, then comparative descriptions often began with apologia for not having seen the types, but they would press on regardless having maybe read about them or seen, at some moment, a painting of one. is kind of doing the best one could in taxonomy was prevalent through the turn of the

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4 20th century and speaks volumes to the perpetual confusion having to do with the age and sex of animals brought in. Without having seen the animals in the wild or living in any way, it was nearly impossible for early taxonomists to accurately describe them. After nearly two paragraphs of both apology and cross reference to what he had read about Pithecia, W. H. Flower in 1862 described a female that came to the British Museum as securely adult, but he confessed that the dimensions of his specimen were rather inferior to either Spixs description of P. hirsuta or Grays of P. irrorata. I examined the very same specimen myself (BMNH No. 1863.6.15.3), and found it to be a small juvenile female.en The age of individuals I had the opportunity to examine available saki skulls in addition to reference skins, mounts, and uids during the course of this revision. And while I only performed the most cursory of examinations (i.e., tenpoint measurements and photographs; see Methods), I noted the relative age-class of individuals based on general tooth wear, canine development, and cranium suturing. I was repeatedly surprised going Table 1. Classications of Pithecia from nominal species to Hershkovitz (1987), Rylands et al. (2000), Groves (2001), and this paper. Nominal Species Hershkovitz (1987) Rylands et al. (2000) Groves (2001) is paper Simia monacha Geofroy St.-Hilaire, 1812 P. monachus monachus P. monachus monachus P. monachus monachus P. monachus P. monachus Geofroy St.-Hilaire, 1812 P. monachus monachus P. monachus monachus P. monachus monachus P. monachus P. hirsuta Spix, 1823 P. monachus monachus P. monachus monachus P. monachus monachus P. hirsuta P. inusta Spix, 1823 P. monachus monachus P. monachus monachus P. monachus monachus P. inusta P. guapo Schinz, 1844 P. monachus monachus P. monachus monachus P. monachus monachus Type not seen P. milleri J.A. Allen, 1914 P. monachus milleri P. monachus milleri P. monachus milleri P. milleri P. napensis Lonnberg, 1938 P. monachus monachus P. monachus napensis P. monachus napensis P. napensis P. monachus Geofroy St.-Hilaire, 1812 P. aequatorialis Hershkovitz, 1987 P. aequatorialis P. aequatorialis P. aequatorialis P. monachus Geofroy St.-Hilaire, 1812 P. isabela sp. nov. P. monachus Geofroy St.-Hilaire, 1812 P. cazuzai sp. nov. P. irrorata Gray, 1842 P. irrorata irrorata P. irrorata irrorata P. irrorata irrorata P. irrorata P. irrorata Gray, 1842 P. irrorata vanzolinii Hershkovitz, 1987 P. irrorata vanzolinii P. irrorata vanzolinii P. vanzolinii P. irrorata Gray, 1842 P. rylandsi sp. nov. P. irrorata Gray, 1842 P. mittermeieri sp. nov P. irrorata Gray, 1842 P. pissinattii sp. nov. P. albicans Gray, 1860 P. albicans P. albicans P. albicans P. albicans Simia pithecia Linnaeus, 1766 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. pithecia Simia leucocephala Audebert, 1797 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. pithecia Simia ruventer Geofroy St.-Hilaire, 1812 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. chrysocephala P. adusta Olfers, 1818 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia Type not seen P. nocturna Olfers, 1818 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. pithecia P. saki Muirhead, 1819 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia Type not seen P. rubarbata Khl, 1820 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia Type not seen P. ochrocephala Khl, 1820 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. pithecia P. capillamentosa Spix, 1823 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. chrysocephala P. pogonias Gray,1842 P. pithecia pithecia P. pithecia pithecia P. pithecia pithecia P. chrysocephala P. chrysocephala I. Geofroy St.-Hilaire, 1850 P. pithecia chrysocephala P. pithecia chrysocephala P. pithecia chrysocephala P. chrysocephala P. monachus lotichiusi Mertens, 1925 P. pithecia chrysocephala P. pithecia chrysocephala P. pithecia chrysocephala P. chrysocephala

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5 back through the data at the disparity in what appeared to be adult skinsbased on size, facial color, pelage, or reproduc tive statusthat they had older juvenile or subadult skulls. is certainly lead to some confusing early taxonomic de terminations, including the fraught P. monachus type in the Musum National dHistoire Naturelle in Paris, which is a long-faded juvenile male based on canine size and body size of the overall mount (cf. P. monachus). Based on skull morphology, some females (among museum specimens) that appeared to have been lactating, or at least were preserved with noticeable, elongated nipples, that may or may not have had distended vulva associated (see Pithecia Females, below), often were juveniles. According to Pereira and Fairbanks (1993), age of reproduction corre lates to juvenile mortality and adult longevity. In primates whose females reproduce younger, there tends to be less juvenile mortality and longer-lived adults. Skins that had signs of reproductive maturation characteristics, such as well-developed throat glands in males or nipples in females, often did not have skulls that were fully adult. It appears that Pithecia may live longer in the wild and have a longer window of reproduction than may have been determined previously, but this clearly needs further investigation.Table 2. Interpretation of Hershkovitz (1987): specimen number call-outs and comparisons with this paper. AM = Adult male, AF = Adult female, JM = juvenile male, JF = juvenile female. Hershkovitz (1987) is rst appearance; many used several times. Museum Specimen No. Hershkovitz ref. Hershkovitz det. LKM det. FMNH 88862 Figure 8 P. m. monachus (AM) P. monachus (AM) FMNH 71806 Figure 8 P. m. monachus (AF) AMNH, not FMNH with that number P. napensis (AF) FMNH 46176 Figure 8 P. p. pithecia (AM) P. pithecia (SAM) AMNH 94132 Figure 8 P. p. pithecia (AF) P. chrysocephala (JM) FMNH 46172 Figure 11 P. monachus (AF) typo FMNH 46176: P. pithecia (M) FMNH 91806 Figure 13 P. monachus (AF) typo AMNH 71806: P. napensis (AF) UCMVZ 157795 Figure 13 P. monachus (AM) P. napensis (SAM) AMNH 93255 Figure 14 P. pithecia (JF) 9325x series in FMNH, but no skull in either AMNH or FMNH with this number FMNH 86995 Figure 15 P. monachus (AF) P. aequatorialis (JF) FMNH 79387 Figure 15 P. pithecia (AM) P. chrysocephala (SAM) FMNH 87000 Figure 16 P. m. monachus (AM) P. isabela sp. nov. (AM) FMNH 86993 Figure 16 P. aequatorialis (SAM) P. aequatorialis (SAM) Art Figure 17 P. monachus unknown amalgam Art Figure 17 P. irrorata irrorata unknown amalgam Art Figure 18 P. monachus unknown amalgam PHOTO Figure 24 P. m. milleri (AF) P. milleri (AF) PHOTO Figure 25 P. i. irrorata (AM) P. mittermeieri sp. nov. (AM) PHOTO Figure 26 P. i. irrorata (AM) P. irrorata (AM) FMNH 122796 Table VI P. monachus (AM) P. isabela sp. nov. (AM) FMNH 70638 Table VI P. monachus (AM) P. milleri (AM) FMNH 79635 Table VI P. monachus (AM) Typo 70635: P. milleri (AM) FMNH 70641 Table VI P. monachus (AM) P. milleri (AM) FMNH 122797 Table VI P. monachus (JF) P. isabela sp. nov. (JF) FMNH 70636 Table VI P. monachus (JF) P. milleri (JM) FMNH 93251 Table VI P. pithecia (AM) P. pithecia (SAM) FMNH 93252 Table VI P. pithecia (AM) P. pithecia (SAM) FMNH 95504 Table VI P. pithecia (AM) P. pithecia (AM) FMNH 95508 Table VI P. pithecia (AM) P. pithecia (JM) FMNH 95509 Table VI P. pithecia (AM) P. pithecia (AM) FMNH 95510 Table VI P. pithecia (AF) P. pithecia (AF) FMNH 93250 Table VI P. pithecia (JF) P. pithecia (JF) FMNH 93253 Table VI P. pithecia (JF) 9325x series in FMNH, but not in either AMNH or FMNH FMNH 95511 Table VI P. pithecia (JF) P. pithecia (JF)

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6 Studies of comparative cranial morphology struggle in that, as with genetics, they tend to mix species together. For in stance, in Marroig and Cheverud (2004, 2009) and Marroig et al. (2003) skulls used for comparison, according to the few reported specimen numbers and location maps, lumped several species. In Figure 2 of their 2004 paper, the P. monachus sample comprised animals in Brazil from Benjamin Constant south along the Ro Yavari into Ro Galvez ( P. monachus), southwest of the Ro Ucayali ( P. inusta), as well as one from northern Peru (no number, but likely P. aequatorialis or P. napensis by location), and one in Colombia ( P. milleri). ey encountered a similar problem with P. irrorata, where none of the animals per the map (or few samples mentioned) are actually P. irrorata (see the species sections in this paper). I observed sagittal cresting in some skulls, but it was not necessarily something that could be used to determine spe cies diferences. It is quite possibly more associated with the kinds of foods available in the region and what is being con sumed rather than a species trait, although some species, like P. monachus, appear more structurally prone to it. Comparisons of skull morphology must be made with care and the understanding that the diferences in age can make a skull in the same species appear to be quite diferent if it truly is an older adult versus a subadult. e key in my opin ion is twofold to understanding saki skulls: 1) e slope of the braincase and face in the articulated skull, particularly as the animal ages, but especially when comparing adults, and 2) the wideness at the back of the mandiblewhile not perfect across all species, the mandible is wide in young sakis and becomes more constricted as the animal ages. In females, again for some but not all species, the older man dibles are wider than their counterpart adult males, which have mandibles that tend to be more constricted. Infants, juveniles, and some subadults seem to have bubble heads, but as the animal matures the braincase narrows and stands up higher, depending on the species. Hershkovitz lamented, Consistent size, and cranial or dental diferences between species of the P. monachus group have not been found (p.410). I suspect this is due not only to the morphological age variation in the skulls, but the fact that he combined several species together in a single taxon. His lamentation is further corroborated in that across museums there were proportionally more juvenile specimens than adults, and far more females than males for most species, except perhaps P. pithecia, which was the closest to equal proportions. Additionally, the older the in dividual, the more likely the coloration matures to portray the true type pattern (see Transitional males below). As seen in older captive animals, chromatic characteristics may change the appearance dramatically as facial and body hair falls out, or is added, depending on the species.en Pitheciaen females Another issue that contributes to errors in Pithecia taxonomy is that females of most species can be quite similar, espe cially as preserved specimens. Many (and in some cases all) females in the museums that I have worked in have been mislabeled as male P. monachus. Hershkovitz (1987) stated the problem accurately: Contributing to the confusion was the failure by all authors to recognize sexual dimor phism in facial pilar pattern. is was further complicated by incorrect sexing, as indicated on collectors skin labels of a large number of the preserved specimens I examined in the various museums (p.409). He went on to say, Failure to recognize sexual dimorphism among sakis has been the greatest stumbling block for sorting the species. e striking similarity between females of all species persuaded some taxonomists to treat all as members of the Pithecia monachus group and even some females of the P. pithecia group as conspecic; irrespective of signicant diferences between the males, that of male Pithecia pithecia excepted (p.415). Hershkovitz himself sufered from this confusion leading to his mass lumping of species. Hershkovitz also admitted that it is really sexual dichromatism more than dimorphism that diferentiates the sexes, if not species. e problem was perpetuated with live animals in captivity up through the 1960s where the males of P. pithecia were often placed with males of P. monachus because monachus was considered the female and pithecia the male (C.P. Groves, pers. comm.). In the current worldwide captive population I suspect that most of the white-faced sakis are likely hybrids; the most common being P. pithecia males (or a hybrid male) housed with a P. chrysocephala female (Fig. 1).Figure 1. A mixed, breeding pair at Elmwood Park Zoo: P. chrysocephala female and P. pithecia male. Photo Elmwood Park Zoo.

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Neotropical Primates 21(1), July 2014 7 Something that is very confusing among the females is their genitalia. I nally understood what I was seeing in numerous specimens after viewing several living females of various species: a swollen vulva resembling male testicles (particularly when dried), and a protruding clitoris, which in museum specimens can resemble a penis (Figs. 2 and 3). e swelling varies among specimens, species, and age groups. Very young juveniles often have the swollen vulvar area even in living individuals, suggesting that age of reproductive readiness is within the juvenile class. Since females can also have distinct throat glands like males and often similar coloration, when coupled in an individual with male-like genitalia and no obvious teats, it is no wonder there is confusion!Figure 2. Examples of female sexual swelling in museum specimens and living sakis: (a) Adult female P chrysocephala (BMNH No. 33.12.6.3); (b) juvenile female P. chrysocephala (MPEG 6971); (c) subadult female P. chrysocephala (MZSP No. 4249); (d) adult female Jamari Saki" (MPEG No. 21934; Appendix II); (e) free-ranging juvenile female P. inusta at Isla de Los Monos, Iquitos; (f) freeranging juvenile female P. aequatorialis at Isla de Los Monos, Iquitos. Photos by L.K. Marsh. a c e b d f

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Neotropical Primates 21(1), July 2014 8 Females can also confuse the casual observer in the wild. In Ecuador, P. napensis lives in smaller family groups of two to eight depending on location, where the main pair often has older ospring or a second adult female living with them (pers. obs.). It was my experience tracking unhab ituated sakis in the wild that often the adult males would do elaborate laps around the forest to distract the threat (observer), while the primary female remained behind on a more obvious branch, often accompanied by another adult or large juvenile female ospring. e females would even tually retreat as well, but not before the untrained observer might dub the pair monachus assuming their identical coloration meant they were male and female of that spe cies. Hershkovitz maintained, Chromatic dierences be tween the sexes are absent or insignicant in P. monachus, irrorata, and albicans (p.415). My experience is dierent: while some species are less derived, such as P. hirsuta and P. albicans, in most cases the sexes are easily separated by appearance. Females placed as male type specimens in clude: P. ruventer/ pogonias (adult female P. chrysocephala, same specimen for both type determinations), P. lotichiusi (adult female P. chrysocephala), and one of the Spix P. hirsuta syntypes is a young juvenile female. Transitional males ere are several specimens throughout the collections misassigned as adult males of a new species, but are actually ju venile or subadult males that are transitional. e reason for this is that many of the saki species, and perhaps all to varying degrees, have dramatic transitional males where the young male has color characteristics of the adult fe males or something else entirely. For example, in P. chrysocephala, the juvenile to subadult males are grey-agouti, have orange bellies, white bangs, and indistinct orange/black facesall resembling an odd looking female rather than an adult male (cf. P. chrysocephala). By comparison, the fully adult males are silky black, lack stippling or brownish/ grey pelage, lack the orange belly, and have solid orangeochraceous facial disks. is has led to intense confusion in museum specimens, especially those whose genitalia were removed or lost during taxidermy, or which are confusing (see Pithecia females above). All species have transitional males to some degree and in some cases females as well. e adaptive signicance of this coloration is not known. Likewise the extent and intensity of the ru in males, while in many cases a species trait, can also vary within species. It seems to change with age, but not always. It would be useful to know whether the color of the ru, in species which tend to have brighter rus, is coincident with breed ing status, age, amount of glandular chemical available, or something else. It would also be interesting to know wheth er any of this kind of passive sexual signaling is at all coin cident with female sexual swelling. A lot more work needs to be done on sexual physiology and behavior in general in the genus. Juvenile males placed as adult types: P. monachus (see Discussion in P. monachus section), P. capillamentosa ( P. chrysocephala) P. nocturna/P. ochracephala (two type de scriptions on same mount, Surinamese P. pithecia).MethodsI use the phylogenetic species concept following Groves (2001, 2004) and Rylands and Mittermeier (2009). Since the designation of subspecies is vague and has been as signed in many cases arbitrarily to describe the diversity of Neotropical primates, I have elected to elevate all Pithecia to full species status until evidence is provided to delineate Figure 3. Examples of juvenile male genitalia: (a) P. chrysocephala (BMNH No. 12.5.11.2), (b) P. monachus pet in Iquitos, (c) freeranging P. aequatorialis at Isla de Los Monos, Iquitos. Photos (a) and (c) by LKM, photo (b) by R. Aquino. a b c

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9 them further (Winston, 1999; Van Roosmalen et al., 2002; Groves, 2004, pers. comm.). In 2001, I rst noticed that the sakis at Tiputini Biodiver sity Station, Ecuador, were diferent (Marsh, 2004), and found that publications about sakis were vague, confusing, or simply repeated information from previous publications (Hill, 1960; Napier and Napier, 1967, 1985; HernndezCamacho and Cooper, 1976; Moynihan, 1976; Hershkov itz, 1979, 1987; Kavanagh, 1983; Wolfheim, 1983; Soini, 1986; Fleagle, 1988; Emmons and Feer, 1990, 1999; Kinzey, 1992, 1997; Groves, 1993; Schneider et al., 1993, 1995; Bodini and Prez-Hernndez, 1995; Burton, 1995; Rowe, 1996; Reid and Engstrom, 1996; Nowak, 1997; Eisenberg and Redford, 1999; de la Torre, 2000; Rylands et al., 2000; Heymann et al., 2002). To make sense of what I was seeing at Tiputini, I conducted eld research throughout Ecuador, visited eld sites and wildlife rehabilitation centers in Ecuador, Peru and Brazil, and went to zoos and primate research centers worldwide to see as many living sakis as possible (Table 3). I also studied collections in 36 museums, where I reviewed 876 skins and uids and 690 skulls in US, European, and South America (Table 4). For each specimen, whether uid, skin or mount, I took numerous digital photo graphs, collected all available information from labels, and eld notebooks if available, and went back to any original documents from the original authors, such as Geofroy St.-Hilaire, Spix, Gray, Lnnberg (see references for full list) to get as accurate a x as possible on some of the more vague type localities. I took twelve photo angles and measured ten aspects of most of the available skulls. I consistently mea sured: braincase length, braincase width, zygomatic arch width, greatest orbital distance, nasal constriction, orbit width and height, greatest muzzle width, mandible length, mandible height and, when possible, canine length. To fully corroborate type and museum data with actual ani mals, I also studied photographs of wild and captive animals throughout South America from a multiplicity of sources, in cluding researchers in situ and travelers in areas where sakis live, reviewed the ISIS database for US and European zoos, and worked with the members of the Pitheciine Action Group as well as established taxonomists (C.P. Groves, A.B. Rylands, and A. Kitchener) to validate my ndings. Every attempt to use pre-existing types and species names before naming a new species was made. Any new names followed the rules of taxo nomic nomenclature (ICZN, 1999; Wilson, 1999). While some research on the genetics of Pithecia has been conducted over the years, I have not used the results here since most of the publications do not have photos or speci men identication of the test subjects, and thus I cannot identify the species used per this revision (but see Martins et al., 1992; Fleck et al., 1999; Voss and Fleck, 2011). Because most of the studies have followed Hershkovitz (1987) and assumed that P. monachus, for example, represented a species that covered the vast majority of the Amazon, I am afraid that (to use a metaphor) apples were compared to elephants and the data may not be entirely meaningful as currently published (see Discussion for more on this topic).Conservation StatusPithecia, like all South American primates, sufer from the efects of region-wide habitat disturbances, including fragmentation of their habitats (Marsh, 2003; Marsh and Chapman, 2013) and hunting. Sakis are hunted through out their range for subsistence, pets, trophies (e.g., as a tourist trade commodity in Ecuador as shrunken heads, pers. obs.), and their tails used as feather dusters or their skins for hats (Mittermeier, 1977, 1991; Peres, 2000, 2001; Bodmer et al., 2001; de oisy et al., 2005; Aquino et al., 2009). Work done by Mittermeier in the 1970s in Suri name detailed the culled biomass and consumption pref erence of Pithecia as compared to the rest of the primate community (Mittermeier, 1977). He discovered that while Pithecia was considered a preferred food by 26.7% of the combined indigenous population, they were rare in the regions where he worked, dicult to hunt, small for the (meat) payof, and proportionally were one of the least hunted of the primates in the country (Mittermeier, 1977). In Ecuador, the Amazonian Quichua do not prefer Pithecia meat, although they will eat it if available. ey say that the meat is toxic, and that it cannot be fed to young children or to dogs as they will become ill or even die (pers. obs.). And while Pithecia, as a group, are hunted throughout their distribution, it is the cumulative efects of all combined human-induced pressures that ultimately will decide each populations conservation status. To date, Pithecia as a genus is considered Least Concern (IUCN 2010), but the IUCN Red List of reatened Species classies only P. pithecia, P. monachus, P. irrorata, P. aequatorialis, and P. albicans, as per Hershkovitz (1987). Pithecia albicans is listed as Vulnerable with populations decreasing, but this was a best-guess estimate by L.K. Marsh in 2007 as part of the IUCN SSC Primate Special ist Group Red-Listing workshop. In actuality, there is very little data on any of the taxa in the wild. Examples are: P. pithecia, which has been studied the longest with the most publications, but in particular by Norconk and asso ciates1, and others (e.g., Mittermeier and Van Roosmalen, 1981; Oliveira et al., 1985; Vi et al., 2001; Riveros and Ferreira, 2001; Lehman et al., 2001; Cunningham and Janson, 2006, 2007); P. chrysocephala (see Rylands, 1992; Setz, 1993; Setz and Gaspar, 1997; Setz et al., 1999; Gilbert and Setz, 2001; Gilbert, 2003); P. napensis (called aequatorialis by DiFiore et al., 2007); P. aequatorialis (see Aquino et al., 2009); P. albicans (see Johns, 1985, 1986, Peres 1993); and a handful of other species (Heymann and Bartecki, 1990; Heymann et al., 2002; Frisoli, 2009; Pal minteri and Peres 2012). 1 http://www.personal.kent.edu/~mnorconk/suriname.html; http://www.personal.kent.edu/~mnorconk/venezuela.html.

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10 Table 3. Living Pithecia in captivity or the wild (not including US zoos) studied for this publication. Status Country Location Site Species Wild Ecuador Yasun Biosphere Reserve 0'32"S, 76'19"W Estacin Cientca Yasun de la Ponticia Universidad Catlica del Ecuador P. napensis Wild Ecuador Yasun Biosphere Reserve 00'05"S, 76'19"W Tiputini Biodiversity Station P. napensis Wild Ecuador Sucumbios 0'34"S, 75'34"W Cuyabeno National Reserve P. milleri Wild Ecuador Yasun Biosphere Reserve 0'09"S, 76'25"W Yarina Tourist Lodge P. napensis Wild Ecuador Yasun Biosphere Reserve 0'54"S, 76'30"W Yuturi Tourist Lodge P. napensis Wild Ecuador Orellana 0'49"S, 76'59" Comuna Pamiwa Kocha P. napensis Wild Ecuador Pastaza 0'49"S, 76'59"W Shiripuno River Lodge/Research Station P. napensis Captive Ecuador Ro Arajuno 1'41"S, 77'16"W AmaZOOnico Rescue Center P. milleri Captive Ecuador Guayllabamba, Quito Zoolgico de Quito N/A Captive Colombia Leticia, Amazonas 3'43"S, 70'23"W Maikuchiga Primate Rescue, Amacayacu National ParkP. hirsuta Wild Brazil Manaus, Amazonas 2'00"S, 60'00"W INPA BDFFP2P. chrysocephala Captive Brazil Manaus, Amazonas Universidade Federal do Amazonas P. rylandsi sp. nov Captive Brazil Rio de Janeiro Jardim Zoolgico do Rio de Janeiro P. mittermeieri sp. nov. (M) P. rylandsi sp. nov. (F) Captive Brazil Rio de Janeiro Centro de Primatologia do Rio de Janeiro P. albicans P. mittermeieri sp. nov. Captive Brazil Sao Paulo Parque Zoolgico do So Paulo P. albicans P. chrysocephala Captive Brazil Ananindeua, Par Centro Nacional de Primatas P. rylandsi sp. nov. Captive Peru Lima Parque de las Leyendas P. inusta Captive Peru Huachipa, Lima Parque Zoolgico de Huachipa P. isabela sp. nov. P. inusta (hybrid) Captive Peru Iquitos Zoolgico de Quistococha N/A Captive Peru Iquitos Proyecto Peruano de Primatologa, Manuel Moro Sommo" Primate Center N/A Freeranging Peru Iquitos Pilpintuwasi Rescue Center P. aequatorialis P. isabela sp. nov. Free-ranging Peru Iquitos, Pto. Indiana Isla de los Monos P. aequatorialis P. monachus Captive France Paris Mnagerie du Jardin des Plantes N/A Captive Switzerland Zrich Zoo Zrich P. pithecia Captive Germany Frankfurt Zoo Frankfurt P. pithecia (M) P. chrysocephala (F) Captive Sweden Djurgrden, Stockholm Skansen P. pithecia Captive Scotland Edinburgh Edinburgh Zoo P. pithecia (M) P. chrysocephala (F) Captive England London London Zoo P. pithecia Photos from Rhett Butler (Flickr) and Xyomara Carretero Biological Dynamics of Forest Fragments Project (BDFFP), conrmed by Brian Lenz, visited by LKM

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11 Table 4. Museums visited or studied for this revision, including facilities that did not have any specimens available, but were checked for completeness. No. of specimens is the total number of skins, mounts, uids, or frozen samples for Pithecia studied in the collection. Museum Location No. of specimens No. of skulls Types Notes on types North America County Museum of Natural History (CMNHLA) Los Angeles, California 3 0 American Museum of Natural History (AMNH) New York City, New York 183 8 1 P. milleri (holotopo-) P. rylandsi sp. nov. (holo-) P. inusta (key) P. monachus (key) P. napensis (key) Allen (1914) Marsh Marsh Marsh Marsh Smithsonian Institution Natural History Division (SMITH) Washington, DC 24* 6 P. aequatorialis (holo-) P. rylandsi sp. nov. (para-) P. isabela sp. nov. (holo-, para-) P. irrorata (key) P. milleri (key) P. monachus (key) P. napensis (key) Hershkovitz (1987) Marsh Marsh Marsh Marsh Marsh Marsh e Field Museum (FMNH) Chicago, Illinois 72 5 1 U.C. Berkeley Museum of Vertebrate Zoology (MVZ) Berkeley, California 6 4 South America Universidad Politcnica Nacional (POLI) Quito, Ecuador 7 3 Museo de Ciencias Naturales Quito, Ecuador 0 0 Universidad Catlica Quito, Ecuador 0 0 Universidad de San Francisco Quito, Ecuador 0 0 Museo Amaznico (AMAZ) Quito, Ecuador 1 0 P. inusta (para-) P. monachus (key) Spix (1823) Marsh Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos (UNSM) Lima, Peru 4^ 1 Museo de Zoologa de Universidad Nacional de la Amazona Peruana (UNAP) Iquitos, Peru 0 3 7 Proyecto Peruano de Primates, R. Aquino Collection Iquitos, Peru 4 0 P. vanzolinii (holo-, para-) P. rylandsi sp. nov. (para-) P. irrorata (key) P. mittermeieri sp. nov. (para-) Hershkovitz (1987) Marsh Marsh Marsh Museu de Zoologia, Universidade de Sao Paulo (MZUSP) So Paulo, Brazil 79 111 P. mittermeieri sp. nov. (para) P. pissinattii sp. nov. (para-) Marsh Marsh Instituto Nacional Pesquisas da Amaznia (INPA) Manaus, Brazil 9 9 P cazuzai sp. nov (holo-, para-) Marsh Museu Paraense Emlio Goeldi (MPEG) Belem, Brazil 102 8 1 P. cazuzai sp nov. (para-) P. irrorata (key) P. mittermeieri sp. nov. (para-) Marsh Marsh Marsh Museu Nacional Rio de Janeiro (MNRJ) Rio de Janeiro, Brazil 57 3 7 Centro Nacional de Primatas (CENP) Belem, Brazil 3 0 Instituto de Investigacin de Recursos Biolgicos Alexander von Humboldt (IAVH) Bogot, Colombia 10 8 P. milleri (key) Marshcontinued on next page

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12 Museum Location No. of specimens No. of skulls Types Notes on types Museo de Historia Natural, Universidad de los Andes (MHNA) Bogot, Colombia 3 3 Instituto de Ciencias Naturales Universidad Nacional de Colombia (ICN) Bogot, Colombia 8 4 Coleccin Manuel RuizGarca, Ponticia Universidad Javeriana Bogot, Colombia 0 11 Europe British Museum of Natural History (BMNH) London, UK 85 8 7 P. irrorata (holo-) P. albicans (holopara-) P. pogonias (holo-) P. hirsuta (key) P. pissinattii sp. nov. (holo-, para-) P. inusta (key) P. monachus (key) P. napensis (key) Gray (1843) Gray (1860) Gray (1842) Marsh Marsh Marsh Marsh Marsh Museum fr Naturkunde (BER) Berlin, Germany 35 1 6 P. albicans (para-) Gray (1860) Zoologische Staatssammlung (ZSM) Munich, Germany 4 2 P. hirsuta (holosyn-) P. capillamentosa (holo-) P. inusta (N/A) Spix (1823) Spix (1823) Spix (1823) Anthropological Institute and Museum (AIM) Zrich, Switzerland 4 1 7 P. pogonias (N/A) Naturmuseum Senckenberg (SEN) Frankfurt, Germany 16 1 4 P. m. lotichiusi (holo-) P. hirsuta/P. monacha (syn-) P. isabela sp. nov. (para) Mertens (1925) N/A Marsh Naturalis Nationaal Natuurhistorisch Museum (LEID) [formerly Rijks Museum van Natuurlijke Historie (RMNH)] Leiden, Holland 35 2 6 P. ochracephala/ P. nocturna (holo-) P. chrysocephala (neo-) P. irrorata (key) Temminck (1863) Marsh Marsh Naturhistoriska Riksmuseet (STOCK) Stockholm, Sweden 54 5 0 P. napensis (holo-) P. irrorata (key) Lnnberg (1938) Marsh Museum National dHistoire Naturelle (MNHN) Paris, France 38 2 1 P. leucocephala (holo-) P. pithecia (neo-) P. monachus (holopara-) P. inusta (neo-) P. mittermeieri sp. nov. (holopara-) Poiteau (1822) Marsh Geofroy (1812) Spix (1823) Marsh Royal Scottish Museum (RSM) Edinburgh, Scotland 4 6 Museo di Storia Naturale di Firenze (MSNF) Florence, Italy 4 0 Naturhistorisches Museum Wien (NHMW) Vienna, Austria 18 1 Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) Bonn, Germany 2 0 Magyar Termszettudomanyi Muzeum Budapest, Hungary 2 0continued on next pageTable 4. continued

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13 Perhaps Collar (1997) said it best: Taxonomy precedes conservation [] without the formal structure of names and an agreed system of usage, there can be no understand ing of what exists to be conserved. Prior to this revision, numerous eld guides, primate species books, and other primate references simply did not have enough informa tion to determine not only what species was in a region, but also what was a male or female. One example of many are the beautiful paintings in de la Torre (2000), one of which was supposed to be a P. aequatorialis male, but in fact is a P. napensis female, and the other of a P. monachus male does not resemble any saki species, anywhere, and appears to be more of an amalgam of what a presumed monachus looks like. Worse, for both scientists working in the region and for tourists, neither P. aequatorialis nor P. monachus occur in Ecuador. us, it is impossible to determine the conservation status of any of the animals identied in this monograph, except perhaps through inference in areas such as Rondnia where there is severe deforestation and exponential human population growth. A great deal of research needs to be done on existing populations, the limits of their distributions, and the human impacts they face before we can con dently report on their status. Whenever possible, I ofer a best guess conservation status, but until we update the Museum Location No. of specimens No. of skulls Types Notes on types Asia Kyoto University Primate Research Institute (KUPRI) Kyoto, Japan 0 3 Total 36 876 690 All Marsh references are to this paper. Key refers to Key Specimens determined by Marsh as important clarications to type. *Does not include uid specimens. ^ e Fleck Lima Collection is at AMNH, accounting for 22 skins and 28 skulls tallied in the AMNH collection. Collection now in Lima at UNSM. N/A = Original type no longer available. All photos and measurements of specimens in Colombian museums by Xyomara Carretero. Data from Museo di Storia Naturale di Firenze provided by Dr. Cecilia Veracini. Data from Naturhistorisches Museum Wien provided by Simon Engelberger and Alice Schuhmacher. Data from UC Berkeley MVZ provided by Francisco Ponutal. Data from Zoologisches Forschungsmuseum Alexander Koenig provided by Dr. Gustav Peters. Data from Magyar Termszettudomanyi Muzeum provided by Dr. Gabor Csorba. Data from KUPRJI provided by Dr. Masanaru Takai.Table 4. continuedIUCN Red List of reatened Species with corrected data from the eld, the status previously posted stands (Version 2010.4. ).NEW TAXONOMIC ARRANGEMENT FOR GENUS PITHECIAFor a full history of genus nomenclature, see Hershkovitz (1979, 1987), Groves (2001), and Rylands and Mittermei er (2009). Abbreviations for all museums referenced in this monograph are in Table 5.Genus Pithecia Desmarest, 1804A thorough reanalysis of the genus Pithecia is presented, including species distribution maps, color illustrations of living species, historic plates, photos of type material (skin and skull), gazetteer of all reference materials (Appendix I), and measurements of type material and photos of living animals. In this revision, there are ve original species, three species elevated from subspecic rank, three historic species rein stated, and ve newly described species. e total number of Pithecia species is 16.

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14 Table 5. Museums and their abbreviations used throughout this publication. Museums and abbreviations Anthropological Institute and Museum, Zurich, Switzerland (AIM) American Museum of Natural History, New York City, New York, USA (AMNH) British Museum of Natural History, London, UK (BMNH) Centro Nacional de Primatas, Belm, Par, Brazil (CENP) Coleccin Manuel Ruiz-Garca, Ponticia Universidad Javeriana, Bogot, Colombia (CPUJ) County Museum of Natural History, Los Angeles, California, USA (CMNH) Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogot, Colombia (ICN) Instituto de Investigacin de Recursos Biolgicos Alexander von Humboldt, Bogot, Colombia (IAVH) Instituto Nacional Pesquisas da Amaznia, Manaus, Brazil (INPA) Kyoto University Primate Research Institute, Kyoto, Japan (KUPRI) Magyar Termszettudomanyi Muzeum, Budapest, Hungary (MTM) Museo Amaznico, Quito, Ecuador (AMAZ) Museo de Historia Natural Andes, Universidad de los Andes, Bogot, Colombia (MHNA) Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima, Peru (UNMSM) Museo de Zoologa de Universidad Nacional de la Amazona Peruana, Iquitos, Peru (UNAP) Museo di Storia Naturale di Firenze, Florence, Italy (MSNF) Museu Paraense Emlio Goeldi, Belm, Par, Brazil (MPEG) Museu de Zoologia Universidade So Paulo, So Paulo, Brazil (MZUSP) Museu Nacional dHistoire Naturelle, Paris (MNHN) Museu Nacional Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ) Museum fur Naturkunde, Berlin, Germany (BER) Nationaal Natuurhistorisch Museum Naturalis (frmr. Rijksmuseum van Natuurlijke Historie), Leiden, e Netherlands (NNMN) Naturhistorisches Museum Wien, Vienna, Austria (NHMW) Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRM) Naturmuseum Senckenberg, Frankfurt, Germany (SEN) Proyecto Peruano de Primates, Rolando Aquino Collection, Iquitos, Peru (AQ) Royal Scottish Museum, Edinburgh, Scotland, UK (RSM) Smithsonian Institution, Natural History Division, Washington, DC, USA (SI) e Field Museum, Chicago, Illinois, USA (FMNH) Universidad Politcnica Nacional, Quito, Ecuador (POLI) University of California, Berkeley Museum of Vertebrate Zoology, Berkeley, California, USA (MVZ) Zoologische Staatssammlung Mnchen, Munich, Germany (ZSM) Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany

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Neotropical Primates 21(1), July 2014 15 WHITE-FACED SAKIPithecia pithecia (Linnaeus, 1766) Description. Males. Fully mature adult males are black with little or no stippling of the dorsal hair. e 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 extend ing from the top of the forehead to the mid-glabella. e 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). Fe males. Varying throughout the distribution, fully mature females are brownish to grayish with some cream or owhite stippling in the dorsal hair, light to bright orange ventral chest hair, and black hands and feet. Females have Synonymy1766 Simia pithecia Linnaeus, Systema Naturae 12th ed. 40. Location unknown. 1789 L Yarke, Buon. Histoire Naturelle Generale et Particuliere, a lHistoire des Animaux Quadrupedes, Suppl. Tome Septime, Paris: LImprimerie Royale. Location unknown. 1797 Simia leucocephala Audebert, Histoire Naturelle des Singes et Makis, 6eme fam., 1: 9, plate 2. French Guiana. 1803 Callithrix pithecia (Le saki ventre roux) Geoffroy Saint-Hilaire. Catalogue des Mammifres du Museum National dHistoire Naturelle, Paris, No. 9. Type no longer exists, female. French Guiana. (see Discussion). 1812 Simia ruventer / Pithecia ruventer Georoy Saint-Hilaire. Recueil dObservations de Zoologie et dAnatomie 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 rubarbata Kuhl. Beitrage zur Zoologie und Vergleichenden Anatomie, p.44. Hershkovitz (1987) notes holotype juvenile mounted in Leiden, juve nile female. Suriname. 1820 Pithecia ochrocephala Khl. 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; in part. Holotype. A type specimen is not known to exist (Hersh kovitz, 1987). Neotype. Simia leucocephala mount No. 452, placed in the Museum dHistoire Naturelle, Paris in 1822 (collector appears to be S. Foiteau) is here designated as a neotype. 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.

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16 distinct white to bufy-orange muzzle lines. e 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 mea surements of adult P. pithecia males and females in the museum collections. Table 7 gives examples of cranial mea surements for adult male and female P. pithecia. Diagnosis. is saki is one of the most familiar of all sakis as it survives well in captivity, even outside of South Amer ica (ISIS 2012, ). Its success in temperate climates may be due to its more northerly distribution. ey are distinct from P. chrysocephala, in that the latter has bright orange to dark ochraceous facial disks, and obvious, stif white to bufy hairs along the lips. Further diferentiation 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. roughout the northern South American countries of Venezuela, Guyana, French Guiana, and Suriname, and in northern Brazil in Roraima, Amap, and parts of Par. ose south of this region are P. chrysocephala. is does not reect 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 dis tinct or are color variations of one or the other species (see Table 6. Average weights and measures for P. pithecia adult males and females in museum collections. In parentheses, the number of specimens/range. Country Male Total body (mm) Female Total body (mm) Male Weight (g) Female Weight (g) Brazil* 767 (11/731) 748 (8/715) 2,057 (3/1720) 1,650 (2/1550) French Guiana 781 (6/747) 768 (4/730) 1,725 (2/1649) N/A Guyana 815 (8/740) 743 (4/715) 2,054 (3/1814-2268) 1,588 (1) Suriname 777 (8/747) 730 (4/712) 1,925 (3/1769) 1,530 (1) Venezuela N/A N/A N/A N/AAll calculations based on available data. Only adults were calculated. Total body is body length including the head and tail. *All locationsTable 7. Skull measurements: examples of adult male and female P. pithecia in French Guiana and Brazil. Measurement (mm) Male* MNHN 1998-233 Female* Male MPEG 21532 Female MPEG 21533 Length of brain case 76.4 73.6 79.5 78.1 Width of brain case 39.7 40.0 47.0 45.0 Zygomatic arch width 53.5 51.3 57.1 51.8 Orbits outer 39.9 41.0 43.9 40.0 Nose bridge 5.0 6.6 6.0 7.9 L-orbit inner width 17.0 16.4 17.3 14.2 L-orbit inner height 16.5 17.5 18.2 14.8 Muzzle width 22.2 19.6 22.8 31.2 L-mandible length 50.0 47.4 51.9 47.9 L-mandible height 31.4 26.6 36.1 33.5 Total body length (mm) 750 800 780 740 Weight (g) 1640 N/A 2500 1750*Male: Museum National dHistoire Naturelle (MNHN), Paris, No. CG 1998-233, French Guiana, Barrage de Petit-Saut, collected by Jean Christophe Vi, 9 Oct. 1994. Female: MNHN No. CG 2001-1981, French Guiana, Association Kwata. Museu Paraense Emlio Goeldi, Nos. 21532 (M) and 21533 (F), Brazil, Par, Oriximin, Cachoeira Porteira, collected by I.E.C. 24 March 1979 and 11 Sept. 1978, respectively. ese are some of the animals in the region where the males have white faces with orange cheeks and cop pery/black pelage, appearing intermediate between P. pithecia and P. chrysocephala, and females more strongly resemble P. chrysocephala.

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17 Venezuela 1. Botanical Garden, Orinoco 2. Lago Guri 3. El Manaco, 59 Km SE of El Dorado at Km 74. Guyana 4. 6.966667, -58.516667 5. Demerara Coast Region 6. Bonnesique Creek (formerly Bonasika River) 7. River Supinaam (by gazetteer Stephens & Traylor 1985) 8. Kartabo (by gazetteer Stephens & Traylor 1985) 9. Kalacoon (by gazetteer Stephens & Traylor 1985) 10. Bartica 11. Rockstone 12. Demerara River 13. Kaieteur Falls 14. Rupununi River Suriname 15. Kapoeri Creek 16. Matapi 17. Between Matapi Creek and Kabalebo River near Corantijn River 18. Kabalebo River 19. Arrawarra Creek 20. Coppename River, Lolopasi, west bank, across from Foengoe Island 21. Voltzberg area east bank Coppename River 22. Brownsberg National Park, M. Norconk study area 23. Saramacca River, Loksie Hattie 24. Bigi Poika 25. Zanderij 26. Hanover, Weg nearby 27. Paramaribo 28. Perica River 29. Wia-Wia Nature Preserve 30. SW of Moengo 31. Moengotapoe 32. Nassau Gebergte near Marowijne River 33. Brownsberg, general area for R. Mittermeier study (1970s) 34. Lely Gebergte 35. Wilhelmina Mountains, approx. West River 36. approx. 1970s Sipaliwini airstrip 37. Kayser Mountains, approx. M. van Roosmalen study area 38. Kayser Gebergte, current airstrip nearby 39. Zuid River 40. Paloemeu River and Tapanahoni River French Guiana 41. Maroni River/Crique Maihonni/ Marowijne River 42. approx. St. Laurent du Maroni 43. Saut Sabbat 44. St. Laurent-Kourou Road 45. Saut Tigre 46. Cayenne 47. Ipousin (by gazetteer Stephens & Traylor 1985) 48. Oiapoque 49. Arouani River Brazil 50. Forte do Rio Branco 51. Rio Cacipor 52. Northern Rio Paru do Oeste, formerly Rio Erepecur (by gazetteer Paynter & Traylor 1991) 53. Serra do Navio Amap 54. Rio Tracajutuba 55. Rio Marac 56. Rio Jari 57. Porteira 58. Rio Paru do Oeste 59. Oriximin 60. Obidos 61. Bravo, Lago Flexal, Edo. do Eira/Lara 62. Boiu (by gazetteer Paynter & Traylor 1991) 63. Patau (by gazetteer Paynter & Traylor 1991) Map 1. Geographic distribution of Pithecia pithecia based on museum specimens and eld study sites.

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18 Discussion). In Brazil, it occurs north of the Amazon and primarily east-northeast of the Rio Trombetas, extend ing 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 in terior 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 Ro Orinoco. eir distribution in the Parque Nacional Canaima is unknown but likely as they are in northern Roraima. eir distribution further west and south in Venezuela is unknown. Specic locations. Appendix I. BRAZIL. Amap: Rio Jary Tapuhy, Cachoeira de Sto. Antnio; Mazago Rio Marac, Moreira, Alto Rio Branco; Terezinha Rio Amu pari, Sinturinha, Teomi, Serra do Navio; Rio Tracajatuba; Rio Villa Nova; Rio Cacipor do Ohcana; Municipality of Oiapoque Villa Velha do Oiapoque. Para : Boiu; Bravo Edo. do Eira; Cabeeiras do Rio Paru do Oeste Posto Tirias; Oriximin Rio Saracazinho, Cachoeira Porteira; bidos Colnia do Veado; Lago Cuitea; Patana; San tarm (captive). Roraima: Forte do Rio Branco; So Joa quim. FRENCH GUIANA. Cayenne: Ipousin Rio Appr ouague. 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: Bartica; Kaieteur Falls; Ka lacoon; 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. ere has been a great deal of confusion over P. ruventer. It was originally included in the Catalogue des Mammifres du Museum National dHistoire Naturelle, Paris, No. 9 (1803) by Geofroy Saint-Hilaire as Cal lithrix pithecia, Le saki ventre roux, without a number. As of 2007 in Museum National dHistoire Naturelle, Paris, there is no type specimen available. ere is, however, a stand with a label on it in the type specimen vaults which reads: #457, Saki a ventre roux, P. ruventer, 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. Geofroy Saint-Hilaires (1851) cata logue, under S[aki] ventre roux, P. ruventer (1851: 55) only one specimen is listed as having been accessed prior to that datea 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 identied in the col lection. e rst use of the name ruventer appears to be by Geofroy Saint-Hilaire in Humboldt and Bonplands 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 Introduc tion). 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. ruventer, Type of Pithecia pogonias Gray (see P. chrysocephala: Discussion). By location alone (French Guiana, per Geofroy Saint-Hilaire), I have added those P. ruventers to P. pithecia. Most of the specimens labeled as P. ruventer throughout the worldwide museum collections are P. chrysocephala females or subadult males from Brazil, or P. pithecia females from Suriname, southern French Guiana, or Amap. 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. ruventer as a nomen oblitum. ere 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 cat egories of P. pithecia and are instead speciesor subspeciesdening characteristics. It appears that further north and west to Venezuela, the females are very simple in color ation: grey stippled, shaggy dorsal body hair, orange bellies, and dark grayish-black, mostly uniform faces with white to bufy muzzle lines. In this region, the adult males are generally very black without stippling in the dorsal hairs, with clean white faces. e juvenile and subadult males can be transitional in that they resemble females in the griz zling of the dorsal hair and bufy 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 bufy to orangish cheeks, or the disk is occasionally entirely light orange in color. Additionally in French Guiana, the ani mals 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 fe males, 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

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Neotropical Primates 21(1), July 2014 19 populations to genetically determine if they are distinct in any way or if these specimens are simply o-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 Figure 4. Likely a painting of the original type, now lost. Origi nal title: Fox-tailed Monkey, Simia pithecia. Plate facing p.169 of Museum Leverianum: Specimens from the Museum of the Late Sir Ashton Lever, by George Shaw. Published by James Parkinson, proprietor, 1792. Figure 5. (a) Adult male neotype of Simia pithecia, and holotype for Simia leucocephala, mount No. 452, Museum dHistoire Na turelle, Paris. (b) Adult male neotype mount, detail of face.a b by exposure. M. Georoy thinks the colour depends on the size of the specimen (p.59). is is another of the taxo nomic confusions early on where they had P. chrysocephala and P. pithecia transitional males standing as types, mixed in with adult males of true species. is reinforces the im portance of seeing these animals in the wild. Even with such a well-studied group such as P. pithecia, we have more work to do.

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Neotropical Primates 21(1), July 2014 20 Figure 7. Juvenile transitional male. Living captive animal, photo by T. Gregory, Suriname (with permission of T. Gregory). Figure 8. Juvenile transitional male mount, Leiden Naturalis No. 39097, type specimen for both Pithecia nocturna and Pithecia ochrocephala, photo by L. K. Marsh. Figure 6. Wild Pithecia pithecia in Venezuela. (a) male, photo by K. E. Glander, and (b) female, photo by M. A. Norconk. Used with permission, all rights retained by Springer Publishers. a b

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Neotropical Primates 21(1), July 2014 21 GOLDEN-FACED SAKIPithecia chrysocephala ( I. Georoy Saint-Hilaire, 1850)Synonymy1823 I. Georoy Saint-Hilaire. Description Pithecia capillamentosa Spix. Simarium et Vespertiliarum Brasiliensis Species Novae [] Le Voyage dans lInterieur du Brasil, Monaco, Species Novae p.16, pl. 11. Holo type registered by Kraft (1983) at ZSM. is same mount had many determinations, including P. ruventer Georoy Saint-Hilaire, P. leucocephala Georoy 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. Georoy Saint-Hilaire. Comptes rendus hebdomadaires des sances de lAcademie de sciences, Paris 31: 875, Brazil. 1850 Pithecia chrysocephala I. Georoy Saint-Hilaire. De scription des Mammifres Nouveaux ou Imparfaite ment Connus de la Collection du Museum dHistoire Naturelle, et Remarques sur la Classication et les Caractres des Mammifres, Second Memoire: Singes Americains. Pp.557, pl. XXIX of type, Brazil. 1852 Pithecia chrysocephala I. Georoy Saint-Hilaire. Catalogue de Primates du Museum dHistoire Na turelle, 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, Manacapur. 1925 Pithecia pithecia lotichiusi Mertens. Senckenbergiana 7(3/4): 74. Brazil, Manacapur. 1939 Pithecia monacha monacha Tate. Bull. Am. Mus. Nat. Hist. 76: 221. Brazil, Amazonas, Jamund. 1987 Pithecia pithecia chrysocephala Hershkovitz. Am. J. Primatol. 12: 417; in part. Holotype. P. chrysocephala plate XXIX in I. Georoy SaintHilaires original Description des Mammifres Nouveaux for the collection at the Museum dHistoire Naturelle, Paris (1850) is accepted here and by Hershkovitz (1987) from I. Georoy Saint-Hilaires 1852 publication. I. Georoy Saint-Hilaire co-types, mounted skinsone male, one undeterminedre portedly acquired in 1850 by the Museum National dHistoire Naturelle, Paris, were not there during Hershkovitzs 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 un known), catalogued on 25 May 1930, acquired from the dealers Schlter & 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 Schlter & Mass in Halle an der Saale, Germany (C. Smeenk, pers. comm.); Manacapur, 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 Manacapur nake Manaos [sic] mittlerer Amazonas, Brazil. Type locality. I. Georoy Saint-Hilaire stated that the ho lotype was from le Brsil, sur les bords du euve des Ama zones. Neotype from Manacapur, 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

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22 around eyes, nose, and chin. Upper lip has thick, stif 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 dened in a disk as they are in adult males. In older ju veniles, this facial hair can be quite shaggy to adpressed. e 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 bufy arms, legs and tail. e agouti pattern is darker and less extreme than subadult males. Hands and feet are black. e 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 bufy 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. eir chest and belly is dull to bright orange. Diagnosis. Pithecia chrysocephala difers 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. ere are similar females in Guyana P. pithecia populations. Measurements. See Table 8. Distribution. Map 2. Only in Brazil, north of the Rio Am azonas, both sides of the Rio Negro, especially in the lower reaches near Manacapur, east to Faro along the Rio Nha mund, 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 I, 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. e boundaries of P. pithecia and P. chrysocephala are unclear further north in the states of Amap, Roraima, and Par, and need further investigation. Specic locations. Appendix I. BRAZIL. Amazonas: Rio Ica; Rio Negro Manaus (Km 165, 170, 190), Apua Bocabau Chueiro/Cuieiras, Acajutuba, Porto Mau, Igarap do Bolivia, Yavanari, Iaunari, Tabocal; Kastuaria Mirim Rio Purus(?); Iranduba; Itacoatiara; Itapiranga; Rio Solimes Manacapur; Rio Itabani nee Atabani; Rio Uruba; Silves; Uatum UHE Balbina, Vista Alegre, So Sebastio; Aniba Igarap Zinho, Rio Angusta; Lago do Serpa; Lago do Canacary. Par : Rio Erepecur Cachoeira do Tronco; Rio Amazonas Faro, Rio Piratucu, Serro do Espelho, Nhamund. Discussion. e 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: Fe males). In preserved skins and mounts this tends to have the appearance of a scrotum and a small penis. Pithecia capillamentosa. e type of P. capillamentosa Spix in the Zoologische Staatssammlung Mnchen, 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 Amap, or even French Guiana as some taxonomists have determined. ere 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 Geof., 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. ruventer Geof, (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 BThard 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 Table 8. Measurements for P. chrysocephala neotype and key speci mens. Adult male and subadult male in Nationaal Natuurhistorisch Museum, Leiden (NNML), and adult female in Naturmu seum Senckenberg, Frankfurt (SEN). All collected originally in Manacapur, Brazil. Measurement Male NN ML 1 845 Sub-adult male NN ML 1 845a Female SEN 6 692 Skull (m m) Length of brain case 73.5 76.9 71.6 Width of brain case 41.3 40.7 40.1 Zygomatic arch width 45.6* 50.1 46.1 Orbits outer 39.0 40.9 38.5 Nose bridge 0.47 0.52 0.50 L-orbit inner width 17.3 16.5 15.7 L-orbit inner height 17.8 16.9 15.4 Muzzle width 19.9 23.0 19.8 L-mandible length 43.5 48.9 44.0 L-mandible height 28.7 32.1 29.4 Po s tcr ania (c m) Head/body 42 47 44 Tail 37 3938*Estimated, right side broken Numbers assigned by LKM; no mount numbers assigned at time of study (November 2007) at NNML.

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23 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 diculty of dis tinguishing 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. e 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, rep resentative of females, with skull, but since the name has not been in use for a valid taxon since 1899, under the pro visions 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. e type of P. lotichiusi in the Naturmuseum Sencken berg, 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. e original determination for this species per the holotype set forth by I. Geofroy was as P. chrysocephala. Hershkovitz (1987) and others considered it to be a subspecies (P. p. chrysocephala) to t into the two-group scheme. I re-elevate it to species as it is distinctly diferent from P. pithecia in phenotype, skull morphology, and range. ere may be hybrid zones throughout the northern range and into Suriname and the Guianas per discussion in the P. pithecia section. e 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 de termine if these animals are hybrids, a separate subspecies, diferent species, or color morphs of chrysocephala. Figure 9. Plate XXIX of Pithecia chrysocephala, adult male holotype per I. Geofroy Saint-Hilaire (1850) in Description des Mammifres Nouveaux ou Imparfaitement Connus de la Collection du Museum dHistoire Naturelle, et Remarques sur la Classication et les Caractres des Mammifres, Second Memoire: Singes Americains.

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24 1. Iaunari 2. Manacapur 3. Acajutuba (approx.) 4. Iranduba 5. Manaus 6. Igarap do Bolivia 7. Rio Apua 8. 105 km north on BR 174 (approx.) 9. Balbina 10. Itacoatiara 11. Lago Serpa 12. Igarap Aniba (approx.) 13. Silves 14. Tabocal 15. Lago Canaari 16. Rio Itabani 17. Rio Uatum 18. So Sebastio do Uatum 19. Rio Piratuc 20. Faro Map 2. Geographic distribution of Pithecia chrysocephala in Brazil.

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Neotropical Primates 21(1), July 2014 25 Figure 10. Hand-colored lithograph from a drawing by Benjamin Waterhouse Hawkins (1807) of "Pithecia pogonias ( P. chrysocephala female) and the head of Pithecia leucocephala", but here depicted as a P. chrysocephala male. Figure 11. Mount of adult male neotype of Pithecia chrysocephala. (1845) at Naturalis Museum, Leiden, with skull (a) and two aspects of the skull (b and c). Photos by L.K. Marsh. a b c

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Neotropical Primates 21(1), July 2014 26 Figure 12. Adult male Pithecia chrysocephala in the wild and captivity. Photos (a and b) by L.C. Marigo and (c) R.A. Mittermeier, all Brazil (photos used with permission). Figure 13. Female Pithecia chrysocephala. (a) subadult, photo by J.M. Ayres, (b) young adult, Manaus, photo by J. White, and (c) adult, So Paulo Zoo, So Paulo, photo by L.K. Marsh (photos used with permission). Figure 14. Transitional, large juvenile to subadult Pithecia chrysocephala, all near Manaus, Brazil. (a) Subadult male, photos by J. C. Serio Silva, and (b) juvenile male, photo by R.A. Mittermeier (photos used with permission). a a b a b c b c

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Neotropical Primates 21(1), July 2014 27 HAIRY SAKIPithecia hirsuta ( Spix, 1823) Synonymy1823 Pithecia hirsuta Spix, J. B., von. Simiarum et Vespertilionum Brasiliensium Species Novae F.S. Hb schmann, Munich, pp.14, pl. 9. Pithecia hirsuta, p.14. Holotype. Adult male mounted specimen with skull intact (No. 19) at the Zoologische Staatssammlung Mnchen (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. e forests between the Rio Negro and the Rio Solimes, Amazonas, Brazil. As for all Spix types a region is reported, not a specic 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 September 1926 (tag note on 27.8.11.23: Topo type 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 Solimes (Tabatinga, on the frontier with Colombia) and Negro, along the Rio Tonantins (possibly near where the original Spix types were collected). BMNH adult fe males were collected in Brazil, Rio I, Lago do Caroira (an oxbow lake on the south side of the river, in the upper reaches, as per Paynter and Traylor [1991]; likely it has a dierent name today). Rolando Aquino collected a subadult male north-east of the Ro Napo, along the left side of the Ro Tamboryacu, northern Peru (now in the UNMSM). Description. e 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 ru. e upper section of the chest from the neck to the ru is obviously bare where the scent gland is positioned. e 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. Deer (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. (is 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 distinc tiveness). 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. e hair around the face is blacker than agouti and not as short and tight against the face as in males. e 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 Spixs measurements of the holotype, and Table 10 for two adult male key specimens in the British Museum of Natural History (BMNH).

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28 Table 9. Original measurements of Pithecia hirsuta holotype from Spix (1823). Morphology Original (in) mm Trunci (body) 1'5 438.2 Capitis (head) 3" 88.9 Facici 1" 38.1 Caudae (tail) 1' 6n" 465.7 Humeri 4" 101.6 Ulnae 3" 82.6 Palmae (hand) 3" 76.2 Femoria 6" 152.4 Tibiae 5" 133.4 Plantae (foot) 4" 108.0 Angulus Facialis 47 Angulus Cerebralis 44 Table 10. Measurements for key specimens of adult male P. hirsuta at the British Museum of Natural History, London. Measurements Male BMNH 27.8.11.23 Male* BMNH 27.3.6.3 Skull (mm) Length of brain case 78.0 Width of brain case 44.4 Zygomatic arch width 56.4 Orbits--outer 43.9 Nose bridge 0.68 L-orbit inner width 17.7 L-orbit inner height 16.9 Muzzle width 24.3 L-mandible length 50.04 L-mandible height Post-crania (mm) Head/body 500 480 Tail 430 410 Hindfoot 120 Ear 31* Skull available, but was not measured.Diagnosis. Pithecia hirsuta is distinct from P. monachus in that it is quite plain, something monachus got the repu tation for probably because of Geofroy Saint-Hilaires 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. e 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 Solimes in Brazil and Peru, north of the Ro Napo in Peru, and south of the Ro Caquet in Colombia (the Rio Japur 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, Co lombia, and probably the Zona Reservada Gepp 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, Pur, and Cahuinar for a total of 1,868,380 ha (Deer, 2004 originally referenced as P. m. monachus). In Brazil, it is found south of the Rio Japur and north of the Rio Solimes-Amazonas, but how far to the Rio Negro in the east they are found is unclear as Spixs type locality is imprecise, suggesting only they are from the swathe be tween the rios Negro and Solimes. Specic locations. Appendix I. BRAZIL. Amazonas: Rio Negro/ Solimes Tabatinga, Rio Tonantins; Rio I Lago do Caroira. COLOMBIA. Amazonas: Ro Igara-Paran La Raicilla Ravine, La Chorrera; Amacayacu National Natural Park; Ro Cahuinari; Bravo Ravine; Ro Cotuhe; San Jose del Encanto. Pur National Park: Cao Mateo, Cao Arapa, Cao Esperanza, Quebradn El Ayo; Curare Los Ingleses Indigenous Reserve: Cao Curare, Cao Agua Blanca, Cao Zumaeta; Camaritagua Indigenous Reserve / Vereda Madroo: Cao El Boliviano. PERU. Loreto: Ro Napo (north) Ro Tamboryacu. Discussion. e 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 Deer (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 Ro Curaray, as part of the expedition that was indeed in the area in previ ous 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.

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29 Map 3. Geographic distribution of Pithecia hirsuta. Peru 1. Ro Tamboryacu 2. Olalla brothers collection area (approx.), Nov. 1925 Colombia 3. Ro Igar-Paran, La Chorrera, 20 km downstream of La Raicilla Ravine (approx.) 4. Ro Cahuinari 5. Amacayacu National Park 6. Ro Cotuhe, Bravo Ravine (approx.) 7. Ro Cara-Paran 8. Ro Cotuhe, Tarapaca 9. 2'S, 69'W Cao Mateo, Pur National Park 10. 2'S, 69'W Cao Arapa, Pur National Park 11. 1'23''S, 69'39''W, Cao Esperanza, Pur National Park 12. 1'S, 69'W, Quebradn EL Ayo, Pur National Park 13. 1'5''S, 69'22''W, Cao Curare, Curare-Los Ingleses Indigenous Reserve 14. 1'36''S, 69 46'02''W, Cao Agua Blanca, Curare-Los Ingleses Indigenous Reserve 15. 1'42''S, 69'58''W, Cao El Boliviano, Camaritagua Indigenous Reserve / Vereda Madroo 16. 1'19''S, 69'04''W, Cao Zumaeta, Curare-Los Ingleses Indigenous Reserve Brazil 17. Rio Tonantins 18. Tonantins 19. Lago do Caruar (location by Hershkovitz 1987) 20. Tabatinga Figure 15. Spix (1823), plate IX, Pithecia hirsuta holotype and corresponding museum mount (ZSM No. 19). Images courtesy R. Kraft, ZSM.

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Neotropical Primates 21(1), July 2014 30 Figure 16. Pithecia hirsuta Spix, Schwarzbrtiger Mnchae, hand-painted plate by Johann Andres Fleischmann (1835) for the Royal Bavarian Art Institute of Private Piloty & Lhle, Munich, and corresponding syntype mount (ZSM No.15). Images courtesy of R. Kraft, ZSM. Figure 17. Adult male key specimen, full body and details, BMNH No. 27.8.11.23. Photos by L. K. Marsh.a b c

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Neotropical Primates 21(1), July 2014 31 Figure 18. Living male and female Pithecia hirsuta from Colombia. (a) adult male, photo by R.A. Mittermeier, (b) adult female pet in Amacayacu National Natural Park, 2009, photo by X. Carretero. Photos used with permission.a b

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32 enMILLERS SAKIPithecia milleri ( J. A. Allen, 1914) 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. Pri matol. 12: 424, in part. Holotype. Adult male in the American Museum of Natu ral 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 Caquet, near Morelia (alt. 700 ft) at the head of the Ro Fragua. Key specimens. Adult male, FMNH No. 70635, collected by P. Hershkovitz, 3 Jan.1952, determination by Hershko vitz as P. m. milleri, Colombia, Caquet, Florencia, Montaita, 400 m. Adult female, IAVH No. 0601, collected by H. Chiriv and J. A. Mora, 8 March 1973 from Colombia, Putumayo, Puerto Leguzamo, Caucaya stream, Miguel Ve lsquez farm, jungle entering by Limoncocha, north-west of Puerto Leguzamo. Note: Breast with milk, infant cap ture. Group of ve individuals in a swamp area. Specimens examined. irty-ve skins and skulls, wild and captive photo references including those observed and pho tographed 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 no ticeably grayer in the face and pelage. Males. As per Allens (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; un derparts thinly haired, foreneck naked; hairs dark brown with whitish tips on the throat and belly and with yellow ish brown tips over the pectoral region; hands yellowish white, feet whitish grizzled with black. Subadult and ju venile 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. Over all 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 cufs of the hindlimbs. e ruf 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 measure ments from Allen (1914).

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33 Distribution. Map 4. In Colombia, Pithecia milleri is distributed from the foothills up to 500 m surrounding Florencia, east to the area of La Macarena, and south of the Ro Caquet to at least Puerto Leguzamo. It is unclear where the boundary with P. hirsuta lies beyond Mecay. According to Deer (2004), it occurs in La Paya National Natural Park (422,000 ha), Colombia. South of the Ro Putumayo it extends into Ecuador north of the Ro Napo, but it is unclear how far it occurs east between the Napo and Putumayo. It is possible since Millers sakis have been recorded in Cuyabeno, Ecuador, that they could be found across the border west into Peru in the Zona Reservada Gepp, north of the Napo. To date, there have been no records of them in that region. Specic locations. Appendix I. COLOMBIA. Caquet: Ro Peneya; Ro Caquet El Inerno Puerto Santander; La Murelia; Florencia Montaita; La Tagua Tres Troncos. Meta: La Macarena National Natural Park. Putumayo: Caucaya Stream Puerto Leguzamo Miguel Velasquez Farm/ Limoncocha; San Antonio Mocoa; Puerto Limn Indig enous Reservation El Picudo Villa Garzn; Ro Mecaya; Caquet River El Hacha Stream (between Putumayo and Caquet). ECUADOR. Sucumbios: Ro Aguarico Lagarto Cocha; Shushundi; Cuyabeno Laguna Garza, Laguna Grande, lodges, Ro Cuyabeno. Discussion. Pithecia milleri was described as a distinct spe cies separate from P monachus by Allen, but Deer (2004) and Hershkovitz (1987) treated it as a subspecies of P. monachus. It is here elevated back to a species. An Olalla speci men, 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: e Ro Lagarto Cocha is a small tributary of the Ro Aguarico, itself a tributary of the Napo. e current international boundary between Ecuador and Peru follows the lower Aguarico and the Lagarto Cocha. [] Carlos Olallas letter stated that it took three days to canoe up the Ro Aguarico to reach the Lagarto Cocha. [] Carlos had nothing to say Table 11. Pithecia milleri holotype AMNH 33876, topotype AMNH 33877, and key specimens FMNH 70635 and IAVH 0601. Measurement Male AMNH 33876 Male* AMNH 33876 Male FMNH 70635 Female** IAVH 0601 Skull (mm) Total length 81.9 82.7 85.7 76.4 Breadth of braincase 43.3 43.7 46.9 41.0 Occipitonasal length 75.7 Basal length 63 Zygomatic breadth 55.5 57 56.9 Orbital breadth 43.0 43.2 42.8 Nasal breadth midorbit 6.3 7.5 5.0 Nasals 16 15 Postorbital breadth 35 L-orbit inner width 16.6 15.8 16.8 L-orbit inner height 17.7 19.6 17.6 Breadth of rostrum at canines 26.4 26.0 25.5 Palatal length 25 Breadth of palate at M1 14 Maxillary tooth rows 19.6 L-mandible length 52.2 61.3 55.6 L-mandible height 41.7 41.8 Post-crania (mm) AMNH 33877 FMNH 70637^ Total length 880 730 819 825 Head-body 480 330 361 420 Tail 400 400 458 405 Hind foot 120 110 120 117.2 Ear 34 31.5* Reported in Allen, J.A. 1914. New South American Monkeys. Bull. Am. Mus. Nat. Hist. 43: 650. Juvenile male topotype does not have its associated skull, only basic body measurements. ^ Adult Male FMNH 70637, same location as key specimen male FMNH 70635, but without skull. FMNH 70635 without body measurements. **Wt. 2102 g. Data on IAVH 0601 from Xyomara Carretero.

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34 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 0'18"S, 75'41"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 Ro Orteguaza appear to be P. milleri, the females, at least in museum col lections, almost resemble P. hirsuta with blacker faces than the more typical P. milleri of Ecuador and southwestern Caquet. As with many saki species, it is possible that the colorations change according to age. us, more research is needed on these populations to determine the boundaries with P. hirsuta. Map 4. Geographic distribution of Pithecia milleri. Colombia 1. La Macarena National Park, Meta 2. Caucay Airport, Puerto Leguzamo 3. Quebrada Caucay, Limoncocha, Miguel Velsquez Farm (approx.) 4. Between Leguzamo and La Tagua (approx., K. Watanabe Expedition 1973) 5. Tres Troncos (approx.) 6. La Tagua 7. Quebrada El Hacha (approx., Caquet River Expedition 1960) 8. Ro Caquet, Puerto Santander, El Inerno Farm 9. Montaita 10. Morelia, Caquet 11. Mocoa 12. Puerto Limn, Villa Garzn, El Picudo Indigenous Reservation 13. La Solita Creek, Solita 14. Vereda La Leona, Valparaiso 15. Ro Peneya Ecuador 16. Ro Aguarico, Boca Lagarto Cocha (by gazetteer, Paynter 1993) 17. Cuyabeno National Park, all points within, including Laguna Garza Cocha 18. Cuyabeno Lodge 19. Ro Aguarico 20. Sushundi

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Neotropical Primates 21(1), July 2014 35 Figure 19. Holotype specimen, adult male, full body and details, AMNH 33876. Photos by L.K. Marsh. Figure 20. Holotype AMNH 33876, skull. Photos by L.K. Marsh.

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Neotropical Primates 21(1), July 2014 36 Figure 22. Adult female Pithecia milleri in Cuyabeno National Park, Ecuador. Photos by L.K. Marsh. Figure 21. Adult male Pithecia milleri in Cuyabeno National Park, Ecuador. Photos by L.K. Marsh.

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37 enMONK SAKIPithecia monachus (. Geofroy Saint-Hilaire, 1812) Synonymy 1812 Simia monacha Geofroy Saint-Hilaire. Rec. Obs. Zool. Anat. Comp. (Humboldt expedition). Vol. 2, p.359. 1813 Pithecia monachus. Geofroy 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. Geofroy 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 from Peru, lower Ro Galvez, eastern side, at Nuevo San Juan. FMNH No. 87002, adult male, skin and skull, C. Kalinowski, January 1957 from Peru, south of the Ro Amazonas, Ro Maniti, east side, at Santa Ceci lia. MPEG No. 1828, adult male, skin and skull, skin and skull, J. Hidasi, November 1960 from Brazil, Rio Javar, Estiro 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, Ro Yavar, no specic site. FMNH No. 88861, adult female, skin and skull, collected by C. Kalinowski, September 1957, from Peru, Ro Yavar-Mirim, no specic location. AQ 04.6.82, juvenile female, skin only, collected by R. Aquino, 4 June 1982 from Peru, upper Ro Orosa on the west side near the conuence of Ro Chontay. BMNH No. 27.3.6.4, adult female, skin and skull, collected by W. Ehrhardt, 10 September 1926 from Brazil, upper Solimes, at Santa Rita, likely present-day Santa Rita do Weil. Type locality. Geofroy Saint-Hilaires (1813) type descrip tion says, Habitele Brsil? (p. 117). Others have placed it with Spixs Pithecia hirsuta north of the Rio Solimes, but the type specimen itself has an indeterminate locality. Specimens examined. Eighty-one specimens, skins, skulls, and photos of living animals from Peru: ros Yavar-Mirim, Lago Preto, Tamashiyacu-Tahuayo by M. Bowler, Quebrada Blanco, Ro Tahuayo by E. W. Heymann, ros Tapiche and Tahuayo by various tourists; in Colombia, Leticia area by Juan Manuel Renjifo Rey and Glenn Perrigo, and cap tives 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. Geofroy Saint-Hilaire (1813) wrote: Pelage vari par grandes taches de brun et de dor: poils bruns en grande partie et des lorigine, et roux-dors vers lestremit: chevelure ray onnante de locciput et aboutissant au vertex (p. 116). He did not provide measurements of this animal, nor an il lustration 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 pre sumably placed by him. e original text from Khl (1820) repeats in Latin basically what Geofroy Saint-Hilaire de scribed, 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, adpres sis. 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,

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38 white-tipped; head and crown of male yellow, of female white (p. 59), which is signicantly closer (but see Discus sion, below). If the type specimen is used as the denitive reference, then the one presently in the Paris vaults is quite diferent 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: e specimen described by Geofroy was young and in very bad shape, which sadly does correctly reect 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. us, 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 dening coloration (previ ous 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. e 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 Javar, Brazil, is labeled male, but it is a juvenile female (Figure 33). It is clear from this study that the holotype has value for histori cal purposes only, not for taxonomic reference. Description. ere 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 par ticular 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. e forearms and across the chest tend to have longer whitish tips, with the chest ruf having dark roots, but light tan to orangish tips. Juvenile and subadult males are more grizzled in general, with the ruf more ob vious and stippling across the arms at chest height make the ruf seem more extensive. e 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. e hands and feet are mottled black and white, growing whiter with age, but the hairs are subtermi nally black. e adult males faces are very brown, and are darker the older they get. e hair is adpressed especially in older animals, with a denite 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. ere are ne white hairs along the upper lip as well. e facial skin of the muzzle is black with ne 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, depend ing on age, have a white head-band with some brown form ing 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. e 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. e 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 brown ish 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. e ruf on females is darker brown or black, with occasionally lighter tips, that are less extensive than in males. ey 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 ap pears more as a band around the face, with some hair lling in toward the muzzle, rather than the face nely 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 individu als. 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 interuvial areas between the rios Solimes, lower to middle Ucayali, and lower Yavar in Peru extending south to at least Saraya cu/Serra do Divisor, and the lower reaches of the rios Javar to Juru in Brazil. In Peru: Ro Tahuayo region including the Reserva Comunal Amishiyacu-Tahuayo and Quebrada Blanco, north to Ro Maniti and Ro Orosa, Ro YavarMirim and Ro Yavar region, including, Quebrada Esperanza, Ro Galvez, north-east to San Fernando, and Leticia/ Tabatinga. Tourist photos and specimens of sakis in the ros Tahuayo and Tapiche are possibly of subadult males or even of P. inusta males. If these animals prove diferent, it

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39 may be because of introgression from P. inusta in the Ro Ucayali watershed coming up from the south (cf. Discussion below). In Brazil: Benjamin Constant, Santa Rita do Weil, Atalaia do Norte, and Estiro do Equador. Specic locations. Appendix I. BRAZIL. Amazonas: Rio Javari Estiro do Equador, Atalaia do Norte; Benjamin Constant Mata Esperana do Municpio, Rio Quixito Seringal, Boa Vista (Gondino), Jos Veiga; Santa Rita; Tabatinga. PERU. Loreto: Ro Yavari; Ro Yavari Mirim Mariscal Ramon, San Fernando, Quebrada Esperanza, Lago Preto; Ro Maniti Maynas, Santa Cecilia; Pebas; Ro Amazonas Orosa, Ro Chontay; Ro Galvez Nuevo San Juan; Ro Tahuayo Quebrada Blanca, Tahuayo Lodge, Tamashiyacu; Ro Tapiche Puerto Punga, San Salvador; Iquitos. Discussion. e Monachus Mess has been snowballing for almost 200 years. I read all available early references regarding the description of monachus, including Geoffroy (1812, 1813), Khl (1820), Spix (1823), Gray (1860, 1870), Elliot (1913), J.A. Allen (1914), Lnnberg (1938), and Tate (1939). All of them were confused and conicting about what Geofroy meant. Spix (1823) simply ignored monachus altogether, clearly naming and referencing hirsuta and inusta, animals that can still be identied today (cf. P. hirsuta and P. 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. Geofroys description of monachus from the Castelnau Expedition, Spixs P. hirsuta, Poeppigs P. guapa and Lessons two P. nocturna (both of which are P. pithecia transitional juvenile males), and his own P. irrorata saying, the [speci men] in the British Museum, gured in the Voyage of the Sulphur, has the face quite bald. is 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 of (cf. P. irrorata). Lnnberg understood the problems best, and tried to give an historic rationale: P. monachus was named in 1812 by Geofroy St. Hilaire ( Ann. Mus. Nat Paris XIX, p. 116). Although the general characteristics of this monkey are in dicated, it has not been suciently clear which of the ex isting races or subspecies is to be considered as the typical one. In consequence of this the specic name quoted has sometimes been used in a rather collective sense, and sev eral 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, Pl. IX) and inusta (i.e., p. 15, Pl. X). Both these subspecies are represented in the present collection, and as it is not proved, whether any of these can be regarded as identi cal with the typical monachus, the names of Spix are used. Like Spix, Lnnberg never attaches any specimens to monachus, instead he accurately supports Spixs clearly dened species (and adding a new one of his own, cf. P. napensis) leaving us to believe that while Geofroy indeed was look ing 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 Ln nberg 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 Hershkovitzs last saki work, he fell prey to the same condition. Hersh kovitz (1987) stated, Consistent size and cranial or dental Table 12. Measurements for P. monachus key specimens. Skull Measurements (mm) AMNH 11132 Male FMNH 87002 Male MPEG 1829 Male FMNH 88862 Male FMNH 88861 Female Length of brain case 81.6 84.8 86.3 86.9 80.0 Width of brain case 36.8 46.0 44.0 48.2 42.6 Zygomatic arch width 53.0 58.0 57.4 56.8 51.0 Orbits-outer 38.9 41.3 46.2 42.1 39.8 Nose bridge 6.1 7.15 7.0 7.2 6.2 L-orbit inner width 17.5 16.2 17.4 15.6 16.1 L-orbit inner height 18.5 19.4 19.3 18.3 18.4 Muzzle width 24.2 25.4 25.5 26.4 22.8 L-mandible length 52.1 59.1 57.6 61.0 56.5 L-mandible height 38.0 39.1 41.0 40.7 38.1 Post-crania (mm) Total 875 8 864 Tail 457 4 404 Hind foot 126 1 118 Ear 33 3 32

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40 diferences between species of the P. monachus group have not been found (p. 410). is problem for him likely arose from the fact that he lumped several diferent species and sex/age classes into not only the monachus group, but into the monachus species itself (cf. Table 2 Introduction). e 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). e animals depicted in it are from Peru. It is of note that tienne Geofroy Saint-Hilaires 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 recog nizable 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; ompson and Norconk, 2011; ompson et al., 2011), but there are no studies on the size of the glandular areas or excretion rate/ Map 5. Geographic distribution of P. monachus. Brazil 1. Estimated location Rio Juru (E. Garbe Expedition 1901) 2. Santa Rita do Weil 3. Benjamin Constant 4. Atalaya do Norte 5. Rio Quixito 6. Estiro do Equador Peru 7. San Fernando 8. Boca Ro Yavari Mirim 9. Quebrada Esperanza (approx.) 10. Ro Orosa 11. Orosa 12. Ro Orosa (R. Aquino collection) 13. 3'S, 72'W 14. Rio Maniti, Ro. Cecilia 15. Pto. Indiana 16. Quebrada Blanco 17. Tahuayo Lodge 18. Ro Tahuayo, Tamashiyacu 19. Ro Yavari Mirim 20. Ro Galvez, Nuevo San Juan 21. Ro Tapiche, San Salvador 22. Boca Rio Punga, Puerto Punga 23. 6'S, 74'W 24. Sarayacu

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Neotropical Primates 21(1), July 2014 41 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 [2002] for A. palliata), it is unclear wheth er 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. e function of both visual and olfactory cues in saki sexual behavior needs further investigation in all species. e deep confusion over P. monachus was not just over its appearance, but its location, as Georoy confusingly gives us Le Brsil? 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 ros 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 Juru in western Amazonia and Acre; alti tudinal 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 informa tion to mean that monachus was also in Colombia (Deer 2004) and worse, throughout Ecuador, Peru, Bolivia, and Brazil, all the way to the Rio Tapajs (BDGEOPRIM 2002). It is no wonder we have such divergence in genetic results, and in any kind of comparative anatomy for proj ects involving P. monachus. With more extensive analysis and considerably more data, it may be shown that P. inusta and P. 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/Yavar/Juru region. e sakis in the lowermost Ucayali, in particular in the ros 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 Yavar/Javar, 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 dierences 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. Figure 23. Pithecia monachus depicted in Plate 3 of Castelnau et al. (1855).

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Neotropical Primates 21(1), July 2014 42 Figure 24. Pithecia monachus holotype MNHN No. 447. Photos by L.K. Marsh. Figure 25. Full mount and detail of MNHN No. 448, subadult female, from Ro Yavar, collected on the Castelnau et Deville Expedition, 1867. Photos by L.K. Marsh.

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Neotropical Primates 21(1), July 2014 43 a (this column)b (this column) Figure 26. Adult female key specimens, full body and details of: (a) Brazilian Pithecia monachus, BMNH 27.3.6.4 from Santa Rita, Solimes, and (b) Peruvian Pithecia monachus, FM NH 88861, adult female from Ro Yavar-Mirim. Photos by L.K. Marsh.

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Neotropical Primates 21(1), July 2014 44 Figure 27. Adult male key specimens, full body and details of: (a) Brazilian Pithecia monachus, MPEG 1828 from Estiro do Equador on the Rio Javar, and (b) Peruvian Pithecia monachus, FMNH 87002, adult male from the Ro Maniti. Photos by L.K. Marsh. a (this column)b (this column)

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45 Figure 28 (ac) Wild adult male Pithecia monachus, (d) captive adult male, Iquitos Zoo. Photos by Mark Bowler.a c d b Figure 29 Subadult male Pithecia monachus (or possibly adult male P. inusta; cf. Discussion) from the Ro Tahuayo region. (a) wild male, photo by Mark Bowler; (b) provisioned male, photo by John Agnew.a b

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46 Figure 31 Diferent ages of transitional males: (a) Living juvenile male Pithecia monachus near Leticia, photo by Juan Manuel Renjifo Rey; (b) pet in Quebrada Blanco, Ro Tahuayo, Peru, photo by U. Bartecki; and (c) pet in Iquitos, Peru, photo by R. Aquino.a b c Figure 30. Subadult male Pithecia monachus sexually gesturing to a female. Photo by Sally Kneidel.

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47 Figure 32. (a) Juvenile female Pithecia monachus in the Tamashiyacu-Tahuayo region, photo by E.W. Heymann; (b) adult female P. monachus (with squirrel monkey), rescue center near Ro Selva, Tabatinga, photo by Glen Perrigo; (c-d) wild P. monachus in Peru, breeding (subadult) female, near Boca Ro Yavar-Mirim, photos by M. Bowler. a b d c

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Neotropical Primates 21(1), July 2014 48 Figure 33. Various aspects of Pithecia monachus, juvenile female, MPEG 30768, from Atalaia do Norte, Brazil.

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49 enBURNISHED SAKIPithecia inusta ( Spix, 1823) Synonymy 1824 Pithecia inusta Spix, J. B. von. Simiarum et Vespertilionum Brasiliensium Species Novae. 1823. F. S. Hb schmann, Munich, p. 15, pl. 10. 1987 Pithecia monachus monachus Hershkovitz, P. Am. J. Primatol. 12: 422, in part. Holotype. Plate only (Figure 1), original mounted speci men 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 collect ed by Castelnau and Deville in 1867 as part of their South American expedition. UNMSM No. 29, skin only, collect ed by Koepcke in 1949, Peru, near Huanuco, Ro Llullapi chis, Estacin Biolgica Panguana. BMNH No. 28.5.2.42, adult male, skin and skull, 18 September 1927, from Peru, upper Ro Ucayali, Chicosa, and No. 28.5.2.43, subadult female, skin and skull, 3 August 1927, from Peru, upper Ro Ucayali, Cumeria. Both collected at 1500 feet above sea level (457.2 m), and on the Godman-omas Expe dition (omas (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. e 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. us, the following description takes into ac count 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. e ruf is darker brown at the base with light tan tips in younger animals, but can become brighter with bufy-tannish or nearly orange tips in older males. e chest is otherwise sparsely covered in black hair. e forearms in males are sparsely covered in a pattern that for some looks like a tri angle up the arm, where half or more is not stippled at all. Wrists tend to be black or without grizzling. e 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 of-white hairs with white, short-haired muzzle lines that are more or less distinct as they blend with the rest of the facial hair. e muzzle is bare, with dark skin, and there are scattered short white hairs along the lips. e 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 (Ro Tahuayo/Ro Tapiche) have whit ish faces with a tan line down the center of the forehead,

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50 over the indent (cf. P. monachus Discussion); whereas, in dividuals in the south appear to be more uniform in their burnished color as they get older. ere 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. Lnnberg (1938) agreed with the original Spixs (1823) de scription of P. inusta saying: e crown and the forehead, the cheeks from below the eyes and the sides of the head are densely covered with short, very stif erect hairs which are directed somewhat forwards, or so on the sides more or less downwards, but are not adpressed. e 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. e dorsal coloring in adult females is very grizzled, often with a light tan colored ruf, and a mostly white to tannish face, similar to males. Forearms tend to be densely covered with white grizzling and white wrist cufs (lacking in the males). Faces of adult females are similar to males, although they are longer overall and whiter in general. ey 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 cov ering 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. e 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. 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. e species most similar to P. inusta is P. monachus, which likely shares part of the northern limits of the range, particularly between the ros Amazonas/Yavar in the north-eastern corner of Peru. Pithecia inusta is clearly Table 13. Original measurements of Pithecia inusta holotype by Spix (1823). Morphology Original (ft/in) mm Trunci (body) 1'4" 406.4 Capitis (head) 3" 76.2 Facici 1" 31.8 Caudae (tail) 1'7" 482.6 Humeri 3" 82.6 Ulnae 3t" 79.4 Palmae (hand) 2" 57.2 Femoria 4t" 104.8 Tibiae 6" 152.4 Plantae (foot) 4" 101.6Table 14. Measurements for Pithecia inusta in Peru and the Brazil, Rio Juru sakis. Skull (mm) BMNH 28.5.2.42 Adult male AMNH 239853 Subadult male NHRM 2219 Adult male NHRM 2375 Subadult male NHRM 2472 Adult female NHRM 2328 Subadult female Length of brain case 86.0 85.0 90.0 84.0 86.0 86.0 Width of brain case 43.0 45.1 47.0 43.0 45.0 44.5 Occipitonasal length 78 72.5 76 76.5 Condylobasal Length 74 65 70 67 Zygomatic arch width 59.3 57.5 61 50 56.5 52 Orbits outer 43.0 37.6 45.5 40 43 40 Least postorbital width 34 34 35 35 Nose bridge 6.4 6.1 L-orbit inner width 15.5 16.5 L-orbit inner height 16.4 18.4 Palatal Length 31 26.5 28.5 25.5 Breadth of palate insideM1 14 13.5 13 11 Muzzle width 25.1 26.7 27.5 22.5 22.5 23.5 L-mandible length 54.6 50.1 L-Mandible height 39.4 Length of maxillary row of cheek teeth 19 20 19 19 Measurements reported by Lnnberg (1938).

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51 distinct from monachus and all other sakis in the ontog eny 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 Ro 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 Ro Urubamba/ Ro Tambor split. ey are in the foothills along the Ro Pachitea below Oxapampa, particularly east of the Pachitea along the Ro Llullapichis, Monte Alegre, Puerto Victoria, and Nevati/Mission. See Discussion below for sakis in the upper Rio Juru region in Brazil and the upper Ro Pachitea in Peru. Population may reach as far north as the ros Tahuayo and Tapiche in the lower Ucayali watershed (cf. Discussion, below). Specic Locations. Appendix I. BRAZIL. Amazonas: Rio Juru Igarap do Gordo, Igarap Grande, Joo Pessoa. Acre : Upper Rio Juru Porongaba, Porto Saids. PERU. Huanaco: Ro Pachitea Monte Alegre, Tabalosos, Pachitea Pasco: Oxapampa Nevati Mission, Puerto Victoria; Ro Pachitea Panguana, Ro Llullapichis, Ro Santiago Loreto: Ro Ucayali Sarayacu; Contamana Cerro Azul; Cume ria; Masisea Tushemo; Chicosa. Ucayali: Ro Ucayali; Ro Pisqui; Ro Urubamaba; Via Utuginia; Ro Inuya; Lagarto. Discussion. Inustus in Latin means burned, and is an apt description of the adult males with their burnished tanwhite faces. Elliot (1913) was the rst 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 Geofroy give location information for either inusta or monachus: e 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 Tonan tins abuent of the Solimes River near Tabatinga (p. 290) (see e 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 origi nal Spix type is of a large juvenile to subadult male, not an adult. Lnnberg (1938) kept the incorrect location place ment, but allowed that P. inusta was indeed a true species based on Spixs type and description. Hershkovitz (1987) and others placed it incorrectly with the P. hirsuta location as described by Elliot (1913). Likewise, there is no specic 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 Ro Amazonas (Castelnau et al., 1855). Specimens from the Rio Juru, Eirunep (Igarap do Gordo and Igarap Grande) in Brazil are included here as part of P. inusta (as per Lnnberg 1938), although further investigation needs to be conducted in this region. In the northern reaches of the Rio Juru 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, ares on their zygo matic arches, and large sagittal crests, which are closer to monachus than inusta skulls except for the heaviness. e older upper Juru adult males tend to have faces that are intermediate between P. inusta and P. monachus. e 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. e 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 Ro Pachitea region, particularly in the upper reaches (Puerto Victoria and south to Nevati/Mission) are diferent 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. ese are possibly diferent 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 provi sionally included in P. inusta: SEN No. 16605, skull only, with the compelling label information of Pithecia monacha Geofr. M hirsuta Spix, F inusta Spix, location W. Brasilien. Get. 1826 Mnchen Mus. gegen Rppells Dubl. Is this Table 15. Pithecia inusta post-cranial measurements for Peru and Brazil, Rio Juru sakis. Post-crania (mm) BMNH 28.5.2.42 Adult Male AMNH 239853 Subadult Male BMNH 28.5.2.43 Subadult Female NHRM 2219 Adult Male NHRM 2375 Subadult Male NHRM 2472 Adult Female NHRM 2328 Subadult Female Head-body 425 401 396 410 370 420 375 Tail 488 305 375 500 465 545 470 Hind foot 121 135/137 121 130 117 132 125 Ear 29 37 33 Measurements reported by Lnnberg (1938).

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52 the skull belonging to Spixs type in ZSM? Is it one of the Rio Juru sakis? And it is interesting especially with Rp pells 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. e skull appears to be subadult and likely female. e sakis in the lowermost Ucayali, in particular in the area of the ros 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 Yavar/Javar, 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 understand ing that there are diferences 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.Figure 34. Pithecia inusta holotype, Plate X from Spix (1823). c

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Neotropical Primates 21(1), July 2014 53 Figure 35. Key specimens of Pithecia inusta: (ab) MNHN No. 449, subadult male, full mount and detail, and (cd) UNMSM No. 54E, adult female, full mount and detail. Photos by L.K. Marsh. c b d Figure 36. Key specimen, UNMSM No. 29, adult male skin only, full body and detail, from the Ro Llullapichis, Peru. Photos by L.K. Marsh. a

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Neotropical Primates 21(1), July 2014 54 Figure 38. Captive (a) adult male and (b) female Pithecia inusta at Leyendas Zoo, Lima, Peru. Photos (a) L.K. Marsh and (b) R.A. Mittermeier. a b Figure 37. Key specimen, Pithecia inusta. BMNH No. 28.5.2.42, adult male, skin and skull. Photos by L.K. Marsh. a

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55 Figure 39. Captive (a) juvenile male and (b) juvenile female Pithecia inusta at Leyendas Zoo, Lima, Peru. Photos by J. Vermeer. Map 6. Geographic distribution of Pithecia inusta, including populations in the Ro Pachitea, Peru and Rio Juru, Brazil. Brazil 1. Rio Juru, Igarap do Gordo 2. Rio Juru, Igarap Grande 3. Rio Juru, formerly Joo Pessoa (presumed; Olalla 1936 Expedition) 4. Porto Saids, Acre 5. Porongaba, Acre Peru 6. Tabalosos 7. Sarayacu, Olalla camp May-June 1927 8. Sarayacu, Olalla Camp Mar-5 May 1927 9. Sarayacu, Olalla camp Aug 1927 10. Bomba/Rio Bomba 11. Cerro Azul? 12. Contamana 13. Ro Pisqui 14. Pucallpa 15. Maisisa 16. Tushemo 17. Boca Ro Pachitea 18. Montealegre (approx.) 19. Ro Llullapichis 20. Ro Llullapichis, Est. Biol. Panguana 21. Ro Utiquinea 22. Puerto Victoria 23. Pto. Pachitea 24. Cumeria 25. Ro Pichis (right bank), Nevati Mission approx. 26. Chicosa 27. Lagarto (approx.) 28. Boca Ro Urubamba 29. Ro Inuya b a

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56 enCAZUZAS SAKIPithecia cazuzai sp. nov. Synonymy 1987 Pithecia irrorata Hershkovitz, P. Am. J. Primatol. l2: 42426, in part. Etymology. e name was selected for a respected, worldrenowned Brazilian primatologist, Dr. Jos de Sousa e SilvaJnior (Cazuza), who has worked many long years at the Museu Paraense Emlio Goeldi in Belm, now as the Cura tor of Mammals, where he continues to contribute signi cantly to South American mammalogy and taxonomy. Holotype. Subadult male, Museu Paraense Emlio Goeldi No. 37127, skin and skull. Collected by J. Urbechi in v rzea, 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 Servio de Estudos e Pesquisas sobre a Febre Amarela. Large juvenile female MNRJ 3312, skin only, collected by C. Lako, 20 Sept1927. Type locality. Holotype. Uarini, margem esquerda mdio rio Solimes, comunidade Barroso Reserva de Desen volvimento Sustentvel Mamirau. Paratypes. Fonte Boa, Rio Solimes, Amazonas. Specimens examined. ree (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 griz zling 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. e chest is bare with sparse black hairs, including a minimal black ruf under the bare throat patch. Belly appears to be without pigmentation in the prepared specimen (sub adult). e arms have brown cufs at the wrists extending in a brownish V down the backs of the hands. On the ankles, there is also a brown cuf ring on top with a less ex tensive brown V down the feet. e hands and feet are of white, largely because they are grizzled with brown/black hairs. e 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 un pigmented or pinkish. White malar lines are likewise sparse and thin, but the hair continues up under the eyes. e hair on the lips is white, stif, and quite thick. Females. e 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. e hands and feet are white with only very slightly brownish cufs on the wrists, without a V onto the backs of the hands, but with a slight V on the feet. e undersides are barely covered in ne black hairs, with a short dark brown ruf with tannish tips. e 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. e white band dips a bit above the eyes making a diamond in the forehead showing of the

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57 bare skin in the middle. e arch peters out below the ears where it is black and white along the sides of the face. e 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. Subadult/juvenile females are slightly transitional in that they are more grizzled dorsally, have a lighter tan/brown ruf, 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. ese distinct sakis difer 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 ruf, and faces with such a difuse 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 ruf. ey share thick lip hairs with P. hirsuta and P. chrysocephala to the north and northeast of the Rio Amazonas, but difer signicantly from both of those species in that they lack the agouti faces of P. hirsuta males and the orange/ochre faces of P. chrysocephala. e 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 Solimes on either side of the Rio Juru at Fonte Boa and Uarini. Discussion. It is never an easy decision to add a new spe cies 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.Table 16. Measurements of the holotype and paratypes: skin and skulls. Measurement Male MPEG 37127 Female MNRJ 21055 Large juv. female MNRJ 3312 Skull (mm) Length of brain case 83.5 82.3 Width of brain case 43.6 43.2 Zygomatic arch width 60.9 52.6 Orbits outer 45.7 43.0 Nose bridge 6.7 6.7 L-orbit inner width 18.6 15.2 L-orbit inner height 18.3 16.4 Muzzle width 26.5 23.3 L-mandible length 52.0 50.6 L-mandible height 36.3 37.8 Post-crania (mm) Head-body 600 350 480 Tail 300 430 490 Hind foot 115 Ear 34 Weight (g) 3200 2750

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58 Map 7. Geographic distribution of Pithecia cazuzai. Brazil 1. Fonte Boa 2. Uarini 3. Mamirau Reserve

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Neotropical Primates 21(1), July 2014 59 Figure 40. Pithecia cazuzai, holotype subadult male MPEG 37127.

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60 Figure 41. Pithecia cazuzai skull of holotype MPEG 37127. Figure 42. Pithecia cazuzai. Photo of a living male (www.tatamazonstock.com).

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Neotropical Primates 21(1), July 2014 61 Figure 44. Pithecia cazuzai. Adult female MNRJ 21055 skull. Figure 43. Pithecia cazuzai adult female MNRJ 21055 full body and detail.

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62 Figure 45. Wild (a) male and (b) female P. cazuzai in the Mamirau Reserve for Sustainable Development, north of the Rio Solimes, Brazil. Photo by Felipe Ennes. a b

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63 enEQUATORIAL SAKIPithecia aequatorialis ( Hershkovitz, 1987) 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. ree 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, Ro 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 ruf. e hands and feet are white. e faces of the males are black with a dense white horseshoe band of hair around face. e band is adpressed in the holotype, but appears much thicker on living animals. ere are two often dis tinct, white patches above each eye that connect to the white band around the face. ey have white malar lines around the muzzle and white lip hairs. Juvenile and infant males have very distinctly white heads with the adult color ation apparent even when very young. Females. Overall pelage grizzled grayish over black produc ing an overall grey look. ey have an orange ruf that can be as distinct as males, even in juveniles, but not as exten sive. 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. e 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 ruf 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 rufs, 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.

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64 Distribution. Map 8. Pithecia aequatorialis is found only in Peru, south of the Ro Napo and south (left side) of the Ro Curaray to the Ro Tigre in the west (right bank). e northernmost border with P. napensis in this region is unknown. It occurs on both sides of the Ro 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 conrm 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. Specic Locations. Appendix I. PERU. Loreto: Puerto In diana; Iquitos; Maynas Ro Nanay Santa Luisa; north of Rio Tacshacurary boca Ro Machete; Ro Napo Ro 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 incor rectly assigned to the Ro Coca, a northern tributary of the Ro Napo in Ecuador. e original label says northern Peru and with further investigation this proved correct. Basslers expedition was in northern Peru, JanuaryDecember 1924, and this specimen was obtained between Guamathra and [S. Q.] Tutapischea, now Tutapisco. is information was obtained from the original expedition records tran scribed by Schwartzer at AMNH (unpubl. data). Aquino et al. (2009) studied P. aequatorialis in north-eastern Peru in the Ro Itaya basin (Yanayacu, Seis Unidos, Nauta, Blanquillo), Ro Tigre basin (Patria Nueva, Posayo, Coconilla), and the Ro Curaray basin (Quebrada Arabela, Sector Curaray). eir observations extend the range of P. aequatorialis in Peru further south-west to between the ros Tigre and Corrientes, and correctly exclude the area to the north between the ros Curaray and Napo. I do not dispute their ndings; however, they did not take any pho tographs or samples of the animals throughout their study making it dicult to corroborate this denitively Pithecia isabela sp. nov. occurs west of the Ro Tigre and south of the Ro Maran, and Pithecia napensis extends all the way south from Ecuador to north of the Ro Maran, and likely occurs also west of the Ro Tigre watershed (cf. P. isabela sp. nov. and P. napensis). e phenotypes of P. aequatorialis have seemingly changed over time as well. If one examines the specimens collected Table 17. Measurements for P. aequatorialis paratypes and topotype with body measurements for holotype. Measurement Male FMNH 86991 Male FMNH 86993 Male FMNH 86996 Female FMNH 86994 Skull (mm) Length of brain case 82.9 86.0 78.1 81.2 Width of brain case 46.9 47.9 45.0 45.8 Zygomatic arch width 57.5 57.7 51.7 54.7 Orbits outer 41.9 42.7 42.6 42.1 Nose bridge 6.5 6.8 8.0 7.6 L-orbit inner width 16.9 18.5 16.8 15.8 L-orbit inner height 18.4 18.1 15.8 18.1 Muzzle width 24.0 24.2 22.1 23.1 L-mandible length 51.7 56.8 47.2 53.2 L-mandible height 33.23 37.6 32.3 36.8 L-canine length 13.3 11.0 10.9 10.8 Post-cranial (mm) Head-body 888 900 844 853 Tail 474 460 450 448 Hind foot 131 124 121 122 Ear 30 32 28 31 Holotype Male* FMNH 86992 Head-body 870 Tail 405 Hind foot 121 Ear 30* Skull measurements not available, cf. Hershkovitz (1987), Tables V, VIII, and XI.

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65 by Kalinowski in the 1950s (and there are no other speci mens 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 of of the facemuch more like the facial rings on P. napensis, only their white coloration is difuse 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 Ka linowski are distinct and will prove to be a separate popula tion if they still exist. Map 8. Geographic distribution of Pithecia aequatorialis. 1. Puerto Indiana 2. Mazan 3. Ro Tutapisco 4. Ro Tacshacurary/Rio Machete boca (approx.). 5. Ro Nanay 6. Reserva Nacional Allpahuayo-Mishana 7. Nauta (approx. collection site near Nauta) 8. Santa Luisa (exact location unknown. Lower Rio Nanay near Rio Maraon; Stephens & Traylor 1983).

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Neotropical Primates 21(1), July 2014 66 Figure 47. Skin, whole body and detail, of adult male holotype Pithecia aequatorialis, FMNH 86992. Figure 46. Vintage plate XXXII, 236 186 mm, of Simia pithecia Linnaeus ( Pithecia aequatorialis) by Jacques de Seve in Schreber (1775), and (b) Pithecia aequatorialis original painting by Zorica Dabich and Philip Hershkovitz, through the courtesy of Field Museum of Natural History. a b

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67 Figure 48. Skull of adult male holotype Pithecia aequatorialis, FMNH 86992.

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68 Figure 50. Captive female Pithecia aequatorialis Photo by M. Bowler. Figure 51. Captive (a) juvenile female and (b) infant/juvenile male Pithecia aequatorialis. Photos by M. Bowler. Figure 49. Captive male P. aequatorialis. Photo by R.A. Mittermeier. a b

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69 enNAPO SAKIPithecia napensis ( Lnnberg, 1938) Synonymy 1938 P. monachus napensis Lnnberg, 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 Lnnberg (1938), the type was collected in Ecuador at Napo river at an altitude of 2000 feet [810 m a.s.l.] which concurs with Hershkovitzs (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 ros Napo and Curaray are in the north-west. Male key speci men AMNH No. 36468, however, is reported from Baeza, Ecuador. at 3000 feet (914 m), which is not far from the upper reaches of the Ro 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, purchased from a native collector. Adult male, FMNH No. 41500, skin and skull, collected on 11 February 1922 by R. Olalla from Ecuador, Pastaza, Ro 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, Ro Copataza, 80 miles south-east of Sa rayacu: 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 Ec uador and Peru, and observed in the wild in Ecuador by L. K. Marsh. Description. Males. e most striking feature in the males is that the crown of the head is densely covered with short white hair. e facial disk of dense hair forms a partial disk of white that extends no further than the ears. e disk fades into white and gray-black stippling at the bottom. e 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. e 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. ey 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. is coloration of the ruf is conspicuous even at a distance (pers. obs). Male dorsal body hair is black and grizzled with white tips. e 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. e hands and feet are white to of-white with hands typically whiter than the feet. e ventrum is mostly naked with sparse grayishblack hair that is light and wispy as it is in most saki spe cies. 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. e dorsum of the females is black and stippled as in males, but appears grayish overall. e hands and feet are whitish to of-white as in males. e ruf on the females is dark brown with light tan to bufy tips. e facial hair is not short and dense forming a ring as it is in males, but is ne and covers the face, growing mostly downward. e

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70 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. e 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 spe cies rank. It was correctly described by Lnnberg (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. aequatorialiswhich is not as bulky in the body, has a far more extensive orange ruf, has a clearly fully white, dense horseshoe ring around the face, and whose juvenile males and females are distinctly diferent than P. napensiswere 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 diferences 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 Ro Napo from Coca in the west into Yasun National Park to the east. e distribution along the Napo further east is not known although likely as populations have been sampled north of the Ro Curaray as far south as the mouth where it meets the Ro Napo. In the west, specimens have been collected in the foothills west of the Ro Coca and west of the Ro Napo in Baeza at 914 m and Estribaciones at 1500 m, south to the ros 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 Ro Curaray at the Ro Napo. One was photo graphed by R. Aquino near the Ecuador border on the north bank of the Ro Curaray and the left bank of the Ro Nashio. If a specimen from Parinari Cao is from Peru near present day Parinari, north of the Ro Maran (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 Ro Maran and west of the Ro Tigre will be classi ed as P. napensis, while those to the south will be grouped with P. isabela sp. nov. However, there are samples collected along the Maran labeled with villages just north that may actually be P. isabela (cf. P. isabela sp. nov.). Sakis col lected at the mouth of Ro Santiago (right bank, along with Table 18. Measurements for Pithecia napensis holotype and key specimens. Measurement RNHM 1921 Male RNHM 1921 Male FMNH 41500 Male MHN 1932-2817 SA Female Skull (mm) Length of brain case 80.0 84 85.3 82.1 Width of brain case 44.2 45 48.4 43.5 Occipito-nasal length 75 Condylobasal length 65 Zygomatic arch width 52.0 52 58.4 57.0 Orbits outer 42.5 43 45.3 43.4 Nose bridge 5.7 7.2 -L-orbit inner width 16.0 16.5 16.5 L-orbit inner height 17.7 18.5 17.2 Muzzle width 21.6 23 25.1 23.9 Maxillary row of cheek teeth length 21 L-mandible length 48.9 43.0 54.5 L-mandible height 35.5 35.1 36.7 Canine length 12.5 Post-crania (mm) Head-body 870 840 Tail 440 478 Hind foot 122 128 Ear 37 38 Measurements reported by Lnnberg (1938).

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Neotropical Primates 21(1), July 2014 71 Puerto Melendez) will be likewise listed as P. napensis until further data are obtained. Specic locations. ECUADOR. Napo: Baeza; Ro Pastaza. Orellana: Estribaciones; Ro Napo; Ro Curaray; Macas; Ro Cononaco; Yarina; Pamiwa Cocha Estiro do Equador south Villa Yuca; Ro Shiripuno; Yasun National Park; Napo Wildlife Center; Tiputini Biodiversity Station; Ro Maccas. Pastaza: Sarayacu; Ro Bobanaza; Montalvo; Ro Copataza. Morona Santiago. PERU. Loreto: Boca Ro Cura ray; Puerto Melendez; Ro Tigrillo; Ro Maran. Amazonas: Ro Santiago; Ro Caterpiza Villa Huambisa. Discussion. Heymann et al. (2002) report on their survey results from the Ro Curaray: We saw [] Pithecia monachus only north of the Curaray [] and P. aequatorialis only south of the Curaray. e two species of Pithecia dier 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). is confusion from Hershkovitz is now corrected with this publication (also see Introduction, Table 2), where north of the Cu raray we have P. napensis, and south P. aequatorialis; P. monachus is far to the east (cf. P. monachus). ere is a mounted specimen in the Museo Amaznico, 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. is male has a diuse white facial ring and less dened 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 Ro Maran, the lower Ro Tigre (left bank), and west to Ro Santiago to deter mine where P. napensis and P. isabela sp. nov. share a border, if they do, north of the Ro Maran. Animals from the Ro Santiago region, including those from the Ro Cater piza (MVZ Nos. 157794, 1557795) appear to be closer to P. isabela sp. nov. than P. napensis, in particular in the females. ey may prove to be dierent species or subspe cies, 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 Ro 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 conuence of the ros Napo and Aguarico near the Peru border. More data needs to be gath ered in this region. ere 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 Ro Santiago (unpubl. Bassler Collection records, AMNH). But that location is another collectors catch-all of species making it unclear where the specimen was actually from.Figure 52. Pithecia napensis holotype skin, whole and detail, RNHM (Stockholm) No. A60, subadult male.

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72 Map 9. Geographic distribution of Pithecia napensis. Peru 1. Near Nashio (R. Aquino sighting) 2. Ro Curaray, Boca Curaray 3. Between ros Curaray and Nashio (R. Aquino sighting) 4. Ro Caterpiza 5. Parinari (unknown if correct location for Parinari Cano, Bluntsali-Peyer Expedition 1913) 6. Puerto Melendez 7. Bazea Ecuador 8. Estribaciones 9. Yarina Eco Lodge 10. Ro Cononaco, Yasun National Park 11. Napo Wildlife Center 12. Tiputini Biodiversity Station, Yasun National Park 13. Yuturi Tourist Lodge 14. Ro Santiago, Boca Ro Santiago 15. Shiripuno Lodge 16. Sarayaku 17. Ro Bobanaza 18. Montalvo 19. Ro Copataza 20. Ro Pastaza 21. Macas

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Neotropical Primates 21(1), July 2014 73 Figure 53. Pithecia napensis holotype skull, RNHM (Stockholm) No. A60, subadult male. Figure 54. Key specimens: (a) AMNH No. 36468, adult male, and (b) adult female (top) No. BMNH 80.5.6.4 and adult male (bottom) No. BMNH 80.5.6.5. a aFigure 55. Pithecia napensis males: (a) wild in the Napo Wildlife Center, Yasun National Park, Ecuador, photo by Carol Foil; and (b) captive in Ecuador, photo by R.A. Mittermeier. a b b

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Neotropical Primates 21(1), July 2014 74 Figure 56. (a) Subadult female Pithecia napensis pet at the Aangu Community village near the Napo Wildlife Center, Ec uador. Photo by Max Feingold and (b) wild large juvenile male Pithecia napensis at Tiputini Biodiversity Station, Ecuador. Photo by L.K. Marsh. a b Figure 57. Juvenile male Pithecia napensis: (a) mounted specimens, NNNM, Leiden collected by C. Buckley, 1880, Ro Copataza, Ecuador, photo by L.K. Marsh, and (b) living rescued juvenile male, photo by B. Junek. a b

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75 enISABELS SAKIPithecia isabela sp. nov. neck is black with little or no stipples. e hands and feet are white with a distinct black V extending from the wrists and ankles down the backs of the hands/feet. e forearms have scant to light stippling, but more than on the hind limbs, which are nearly without stippling. e tail is lightly stippled in the manner of the dorsum. e facial disk is dense and dark agouti black to dark brown, with scant white throughout. e eyespots are long, more like stripes, and difuse with a distinct line between them ex tending to the dark facial disk. ere are white malar lines with some white hairs covering the nose. e skin of the face is black. e ruf 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. e pelts of some speci mens appear coppery almost shiny. e dorsal hairs are black with light, difuse, short stippling on the back and forearms, very little on the hind legs. e ventrum is sparsely covered with black hairs, the short ruf 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. e hands and feet are white to of-white. White malar lines trace either side of the muzzle. e skin of the face is black, in both, dried specimens and living animals. Synonymy 1987 P. monachus monachus in Hershkovitz, P. Am. J. Pri matol. 12: 422. 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. e 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 cartog rapher on Charles Marie La Condamines historic expedi tion 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 Ro 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 ros Maraon and Amazon. She eventually reached her husband in French Guiana, where they had a happy reunion. Isabels story of courage and strength is an amazing piece of South American history that absolutely needed proper recognition. Type locality. e holotype is from Peru, Loreto, along the Ro Samiria at Santa Elena (130 m above sea level). e paratypes are all from the region of the Ro Samiria with FMNH Nos. 122798 and 122797 in the Reserva Nacio nal 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 Ro Samiria and in the Reserva Nacional de Pacaya-Samiria. Description of male holotype. Overall the pelage is black with very light, short white stippling. e hood that forms in all sakis with the whirl of hair at the base of the

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76 e facial ring is dark black in adult males and agouti brown in young males, with some light white sprinkled through out 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. e 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). e 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 difuse, 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 griz zling throughout. ere is slightly more grizzling in females than males, but distinctly less than in neighboring P. napensis or P. aequatorialis. e ruf is black with dark brown tips, also right under the throat, but females have a less distinct scent gland. e muzzle lines are thick and distinct and the facial skin is black. e facial disk is black and silky in young females, tighter and black-agouti brown in older fe males. Adults can have white possible in the facial disk, and with tiny white eyebrows and a small white star just above the eyes. e 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 difers consistently in the amount of white on the fore headwhere napensis is clearly dense and white on the crown and grizzled white throughout the facial disk, P. isabela is vastly more difuse, 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 ruf, whereas P. isabela has a shorter, darker brown-orange ruf that is occasionally a brighter shade. e males in particular are far less grizzled overall in P. isabela, and, in some cases, the grizzling ap pears nearly absent. Pithecia isabela obviously difers from P. aequatorialis to the north of the Ro Maran in that males have dense, fully white half-circles around their faces with extensive orange rufs on their chests. Juvenile and subadult males are distinctly diferent 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 difuse white haloing along the crown. ere is no obvious ruf color in the young animals and the griz zling across the back is nearly absent. Measurements. See Table 19. Distribution. Map 19. e species occurs only in Peru. e paratypes and most other specimens are from the PacayaSamiria region. e paratype was collected along the Ro Samiria at Santa Elena. Additional specimens were col lected similarly along the Ro Samiria at the Base Atun and Estacin Biolgica Pithecia. Mark Bowler reported the species along the Ro Yanayacu. Tourist photos show them Table 19. Measurements for P. Isabela, including holotype and paratypes. Measurement FMNH 87001 Male FMNH 122796 Male SEN 5304 Male FMNH 86997 SAM FMNH 86998 SAM FMNH 86999 Female FMNH 87000 SAM Skull (mm) Length of brain case 79.7 79.0 83.7 84.1 79.2 80.3 78.8 Width of brain case 44.1 45.1 43.1 47.6 46.1 44.0 42.8 Zygomatic arch width 57.1 57.9 54.3 59.1 38.6 54.6 50.0 Orbits outer 37.8 42.4 45.5 42.1 49.8 42.7 38.3 Nose bridge 6.5 7.3 0.72 7.5 5.9 6.8 5.9 L-orbit inner width 16.9 17.1 16.4 18.1 17.8 16.7 17.7 L-orbit inner height 17.8 17.4 17.3 19.4 17.8 17.6 16.9 Muzzle width 23.4 24.5 23.8 24.4 21.4 24.4 21.5 L-mandible length 50.9 49.9 52.0 53.3 44.9 50.8 49.8 L-mandible height 37.1 34.2 37.5 34.4 31.2 34.8 33.4 L-Canine Length 12.4 11.6 9.9 10.6 11.3 9.4 Post crania (mm) Total body 920 765 862 796 853 867 Tail 486 400 450 433 445 448 Hind foot 131 121 122 122 124 124 Ear 30 33 31 30 30 30

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Neotropical Primates 21(1), July 2014 77 at Quebrada Sapote near the Ucayali, and collections by R. Olalla suggest they are near Sarayacu on the west side. Specic distribution. Appendix I. PERU. Loreto: Ro Samiria Santa Elena, Biological Station Pithecia, Base Atun; Pacaya-Samiria National Park; Ro Yanayacu; Ro Ucayali Sarayacu. Discussion. is species was originally recognized as distinct and suggested as a new species in 2002 at the Tipu tini 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. e 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. ere 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 Ro Maran. It is unclear if these animals ever leave the facilities they were captured for, but if they do, there may be some interesting ramications 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. Bowlers photos were along the Ro Yanayacu, very near to the Rio Maran. It will be important to study animals throughout the PacayaSamiria to determine whether or not P. napensis has in fact established on both sides of the Ro Maraon, and where the border with P. isabela lies.Figure 58. Pithecia isabela. Holotype adult male, skin, FMNH 8700 I.

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78 Map 10. Geographic distribution of Pithecia isabela. Peru 1. Ro Samiria, Santa Elena 2. Yanayacu 3. Nauta -right bank 4. Ro Samiria, Biological Station Pithecia 5. Ro Samiria, Base Atun 6. Reserva Nacional Pacaya Samiria 7. Quebrada Sapote 8. Sarayacu left bank

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Neotropical Primates 21(1), July 2014 79 Figure 59. Pithecia isabela. Holotype adult male, skull, FMNH 87001.

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Neotropical Primates 21(1), July 2014 80 Figure 60. Pithecia isabela. Adult female, skin, FMNH No. 86999.

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81 Figure 62. Juvenile male Pithecia isabela, Reserva Nacional Pacaya-Samiria, Peru. Photo by Rick omas. Figure 61. Wild adult male Pithecia isabela in the Reserva Nacional Pacaya-Samiria: (a) photo by J. Vermeer, (b) photo by Wilderness Classroom.com. a b

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82 Figure 63. Adult female Pithecia isabela: (a) Ro Yanayacu, photo by M. Bowler, and (b) Reserva Nacional Pacaya-Samiria, photo compilation by TrekNature. a b

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83 enBUFFY SAKIPithecia albicans ( Gray, 1860) Synonymy 1860 Pithecia albicans Gray, J. E. Proc. Zool. Soc. Lond. p.231, pl.81. 1863 Pithecia albicans Bates, H. W. e 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 Erhardts expedition are in the Museum fur Naturkunde, Berlin (Nos. 35768, 35769, 35770, 35308, 35767, 46146). Type localities. Holotype: Rio Tef Tef Lake, south bank of the Rio Solim es, Amazonas, Brazil. Syntypes: Amazo nas, Rio Solim es, Rio Purus, Ayapu 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 observa tion of a captive specimen by L.K. Marsh (S o Paulo Zoo, So Paulo). Description. ese 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. e back is black extending from the nape of the neck to the tip of the tail. e tail is entirely black. e wrists often have black short hairs, the hands are of-white, and the feet of-white. Adult males and females have large throat scent-gland patches ringed in a light orange ruf that is more pronounced in males. Males. e facial skin is mostly black, with small bare pinkish skin patches above the eyes, distinct white to cream eyebrows, and stif white to cream upper lip hairs. e muzzle has dispersed whitish hairs, not in a distinct line. Ventrally, adult males have testicles that are light cream to bufy, often with black speckles, and on their kneecaps, what appear to be calluses (obvious on intact specimens both living and stufed, Fig. 70). Females. e 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 sub adults, and may include a short ring of white to of-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. ey share some of the bufy characteristics of P. vanzolinii, but as Hershkovitz (1987) notes, the diferences 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.

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84 Distribution. Map 11. Pithecia albicans occurs only in Brazil. Hershkovitz (1987) places it between the lower Rio Purus and Rio Tef, south bank tributaries of the upper Rio Solimes-Amazonas, Amazonas, Brazil. Per the specimen localities studied for this revision, this distribution is ac cepted specically noting collections near Lago Tef, Lago Ayapu, Lago de Boia, and Jabur on the left bank of the Rio Purus. Johns (1985, 1986) observed P. albicans in terra rma forest on both sides of the Rio Tef, and principally at the Ponta da Castanha (3'S, 64'W) on the eastern side of the lake. ey also occur along the Rio Bauana, a tributary entering the lake on its eastern shore (A.B. Ry lands, pers. comm. in Johns 1986). Peres (1997) observed P. albicans west of the Rio Tef, at a locality called Vai Quem Quer, in terra rma on the right bank of the lower Ro Juru, 3'S, 66'W. Specic locations. Appendix I. BRAZIL. Amazonas: Rio Purus Lago Ayapu, Ayapu town, Jabur; Rio Tef; Tef town; Lago de Boia; Lago Tef Ponta da Castanha.Table 20. General measurements for syntypes (BMNH, London) and other specimens of Pithecia albicans from NHRM, Stockholm. All measurements in mm. Specimen Location Age/sex Total Tail Hind BMNH 27.8.11.17 Lago Ayapu M 920 430 BMNH 34.6.16.10 Lago Ayapu F 880 BMNH 26.5.5.6 Lago Ayapu F 980 460 BMNH 26.5.5.5 Lago Ayapu F 1050 530 BMNH 26.5.5.3 Lago Ayapu M 1030 460 BMNH 26.5.5.4 Lago Ayapu F 1030 490 BMNH 26.5.5.2 Lago Ayapu M 930 420 BMNH 27.8.11.18 Lago Ayapu M 860 400 BMNH 26.5.5.1 Lago Ayapu M 1070 510 NHRM A62-0158 Lago Ayapu F 460 420 103 NHRM A62-0178 Lago Ayapu M 510 500 132 NHRM A62-0293 Lago Ayapu F 990 490 NHRM A63-0169 Lago Ayapu M 890 NHRM A62-1202 Jabur Juvenile F 795 405 120 NHRM A63-1205 Jabur Subadult M 820 416 120 NHRM A62-1207 Jabur Subadult F 810 440 115 NHRM A62-1213 Jabur Subadult F 820 415 120 NHRM A62-1215 Jabur Juvenile F 805 440 100 NHRM A62-1236 Jabur M 850 455 125 NHRM A62-1240 Jabur Juvenile F 820 455 125 NHRM A63-1212 Jabur F 800 415 110* All locations are: Brazil, Amazonas, west of the Rio Purus, south of the Rio Solimes.Table 21. Skull measurements of example specimens of Pithecia albicans from NHRM, Stockholm. Skull Measurements (mm) Male NHRM A62-0178 Male NHRM A62-1236 Subadult female NHRM A62-1240 Total length 84.3 81.1 80.9 Breadth of braincase 43.9 42.6 42.8 Zygomatic breadth 56.6 53.9 53.7 Orbital breadth 45.0 43.9 43.0 Nasal breadth mid-orbit 0.71 0.58 0.75 L-orbit inner width 17.0 17.4 18.2 L-orbit inner height 16.4 17.7 18.4 Breadth of rostrum at canines 24.8 24.3 23.4 L-mandible length 53.0 51.7 48.3 L-mandible height 44.1 38.0 36.6

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85 Discussion. Hershkovitz (1987) and others use the term bufy to mean white, of-white, cream, and tan. e de nition used here and throughout this text for bufy is based on the original tanned hide denition of yellow brown or blonde. us, the bufy 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 Juru), 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 re semblance of P. irrorata and P. albicans has anything to do with a long ago cross-breeding event or if this extremely diferent color variation was just a uke of genetics. In 2008, Pithecia albicans was classied as Vulnerable (VU) on the IUCN Red List of reatened Species (Viega et al., 2008). In 1993, Peres reported the population of P. albicans in the upper Rio Uruc to be 8.8 individuals/km. In 2005, Haugaasen and Peres reported an average of 2.2 indi viduals/km in the lower Purus watershed at Lago Uaua with slightly higher results for terra rma (3.3 individuals/ km) and igap (4.6 individiuals/km), but lower for vrzea (0.8 individuals/km). By all accounts, the population is low if not drastically declining. Research is needed for all areas where P. albicans occurs.Figure 64. Original plate LXXXI of Pithecia albicans from Gray (1860).

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86 Map 11. Geographic distribution of Pithecia albicans. 1. Lago Tef 2. Ponta da Castanha (approx.) 3. Rio Tef 4. Lago de Boia (approx.) 5. Rio Uru 6. Jabur (approx.) 7. Lago Uaua (approx. location of Coari) 8. Lago Ayapu

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Neotropical Primates 21(1), July 2014 87 Figure 65. Pithecia albicans. Holotype skin, adult male, BMNH 60.4.16.3. Photos by L.K. Marsh.

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Neotropical Primates 21(1), July 2014 88 Figure 66. Pithecia albicans. Holotype BMNH 60.4.16.3 skull. Photos by L.K. Marsh. Figure 67. Pithecia albicans. Adult female syntype BMNH 26.5.5.4. Photos by L.K. Marsh. Figure 68. Captive adult male Pithecia albicans. Photo by R.A. Mittermeier.

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89 Figure 69 Pithecia albicans juvenile female pet in the town of Tef, Amazonas State, Brazil. Photo by L.C. Marigo.

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Neotropical Primates 21(1), July 2014 90 Figure 70. Pithecia albicans, adult male underside, showing calloused knee caps, BMNH No. 26.5.5.1. Photo by L.K. Marsh.

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91 enGRAYS BALD FACED SAKIPithecia irrorata ( Gray, 1842) Synonymy 1842 Pithecia irrorata Gray, J. E. Ann. Mag. Nat. Hist., 1st series, 10: 256. 1844 Gray, J. E. e 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, for further synonymy. 2013 Pithecia hirsuta. Van Roosmalen, M. G. M. and Van Roosmalen, T, 2013. On the Origin of Allopatric Pri mate Species and the Principle of Metachromic Bleach ing. E-book, printed privately, p.63. Holotype. Hershkovitz (1987) used Grays (1843) descrip tion from vol. 1, color plate 3. No precise specimen was identied, 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, Ro Manu, Altamira, 350 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 dicult 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 (Payn ter and Traylor 1991). NHRM No. 1153, large juvenile female, skin and skull, collected by A. Olalla, 20 October 1935 from Brazil, Rio Purus, Arum. 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 presentwhite tips of the nape whorlbut it is not dense as in Pithecia rylandsi sp. nov. In subadults, this line seems to trace down the sides toward the shoulders. e 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 Ro Tahuamanu. e hands and feet are white. e faces are pink or un pigmented with a dark muzzle outlined by noticeable, but ne, 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 Ro Manu/Ro Madre

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92 de Dios conuence (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. e sakis further west as far as the Ro 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 ruf rang ing from bright orange to dull orange/light tan. Juvenile males can be browner, with grizzling throughout the dorsal pelage. eir 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). ere 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 bufy as in P. vanzolinii. ey are browner across the front of the shoulders and chest, but do not have a dened light ruf. 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). e 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 difers signicantly from the other bare-faced sakis in the region: P. vanzolinii and P. rylandsi sp. nov. Pithecia irrorata lacks the obvious bufy 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 rufs, something P. rylandsi sp. nov. lacks, but P. vanzolinii loosely shares this feature in that it has a bufy-orange ruf to go with its bufy forearms. And while P. irrorata males in the wild may appear to have dark faces, they are mostly un pigmented except for their black muzzle. Pithecia irrorata females difer 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.Table 22. Measurements for male Pithecia irrorata paratypes FMNH 98040, FMNH 93534 and female paratypes MNRJ No. 3317 and NHRM A62-1153. Measurement FMNH 98040 Male FMNH 93534* Male MNRJ 3317 Female NHRM A62-1153 Female Skull (mm) Length of brain case 85.14 79.99 83.3 77.1 Width of brain case 45.97 48.51 42.5 44.3 Zygomatic arch width 56.88 61.63 54.3 51.8 Orbits-outer 45.90 44.75 45.9 39.2 (est.) Nose bridge 6.88 7.95 8.5 (est.) 0.78 L-orbit inner width 18.51 16.00 16.7 18.4 L-orbit inner height 18.71 17.84 17.0 18.2 Muzzle width 24.38 27.97 25.7 23.2 L-mandible length 59.80 63.19 45.8 48.8 L-mandible height 39.49 41.63 31.3 32.8 Canine length 11.44 13.87 Post-crania (mm) Head-body 419 440 490 440 Tail 496 455 480 407 Hind foot 131 127 130 101 Ear 32 37 18 *Top of braincase missing. Occiput and bottom missing, nose broken on right side. ^ Right orbit broken.

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93 Distribution. Map 12. In Brazil, P. irrorata follows the west side of the Rio Purus, south to Acre. Specically there are specimens from Lago Mapiocu (likely Lago Mapixi) and Arum 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 Juru and Purus, there are not enough specimens to support this, especially as the southern reaches of upper Juru are where P. vanzolinii is found, and the north-west pocket between these rivers is where P. albicans can be found around Tef and, as of 1935, at Jabur on the Rio Purus. It is unknown where the boundaries lie with P. vanzolinii in the upper Rio Juru 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 Rondnia (see Discussion). In Peru, they are primarily in the Manu region, west of the Ro Manu and south of the upper Ro Madre de Dios, west of the Manu conuence. ey extend to the lower Ro Urubamba and are likely to occur in all areas north-east of the ros 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 Ro 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.). eir 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. Specic locations. Appendix I. PERU. Ucayali: Ro Urubamba. Madre de Dios Altamira; Ro Tahuamanu San Lorenzo. BRAZIL. Amazonas: Rio Purus Lago do Arum, Lago do Mapiocu/Mapix Acre : Xapuri; Rio Branco Fazenda Campo Lindo. Discussion. Hershkovitz (1987) describes the history of the type specimen thus: e formal description of P. irrorata appeared in Grays 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, Grays 1842 reference to P. irrorata was evidently in anticipation of his 1843 report [Gray, 1843b]. (p.409). In the spe cies 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 Tapajs, Parque Nacional da Amaznia, Par, Brazil. (p.426). However, animals in this area do not resemble the type as closely as the sakis further south. Hershkovitz found no ref erence 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 nd 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 Hershkovitzs 1987 paper, Pithecia irrorata was not clearly dened (except by implication in his dichotomous key, p. 416), and he went straight to the level of subspecies de notation 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, subspe cies, or diferent species as they deviate from type. ere is much work to be done on P. irrorata throughout its range. e animals in the state of Acre appear to have more grizzling in their overall pelage than those in Peru, es pecially 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. Comisso Rondon sakis. Adding to the mystery of irrorata in Brazil, particularly in Rondnia and Mato Grosso, are col lections from the Comisso Rondon (1909), collect ed largely throughout the Rio Aripuan/Rio RooseveltRio 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 thereexcept per haps 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.).

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94 Furthermore, it is interesting to note that there are some legacy specimens collected as part of the Comisso Rondon (in 1912) that are at present rolled into P. irrorata. ese are a juvenile female, skin and skull (MNRJ 3322) and adult male, skin and skull (MNRJ 3339) both from So 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 Jacaracanga highway. It is en tirely 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 popu lations in the region. Sakis from Jacaracanga. Sakis collected in the early 1970s immediately after the construction of the BR 230 TransAmazonica highway between Itaituba and Jacaracanga on the Ro Tapajs have some interesting variationswhich could be due to them being a diferent 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 diferent. 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 rufand it has a completely pink face! On km 212, col lections on 29 November 1973, by an unknown collector are nearly identical to the km-19 animalsMPEG 8149 an other adult malethis one showing some ne 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. ere 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, 461919both 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 Tapaj s. While is it entirely possible that sakis east of the Rio Tapajs are not related at all to P. irrorata or P. mittermeieri sp. nov., one of the two specimens is similar to the Tapajs females: MZSP No. 3632, subadult female, skin and skull, from Fazenda Marau, San tarm (according to E. Garbe in 1920 this was about four hours away from Santarm, 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 Jacaracanga 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. Figure 71. Pithecia irrorata. J. E. Grays presumed type, from e Zoology of the Voyage of the HMS Sulphur During the Years 1836, Mammalia Plate III, by R.B. Hinds (1843), and as referenced by Hershkovitz (1987) as the nominal holotype.

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95 Map 12. Geographic distribution of Pithecia irrorata. Peru 1. Rio Urubamba 2. lower Rio Urubamba (T. Gregory Field Site) 3. Parque Nacional Manu 4. Cocha Cashu 5. Altamira, RIo Manu (location by gazetteer, Stevens & Traylor 1983) 6. Rio Manu 7. San Lorenzo Brazil 8. Xapuri Acre 9. Rio Branco Acre 10. Rio Purus, Lago Mapixi 11. Reserva de Desenvolvimento Sustentvel Piagau-Purus 12. Rio Purus Aruma (location by gazetteer, Paynter & Traylor 1991)

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Neotropical Primates 21(1), July 2014 96 Figure 72. Pithecia irrorata. BMNH London type No. 101a, skin.

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Neotropical Primates 21(1), July 2014 97 Figure 73. Pithecia irrorata. BMNH London type No. 101a, skull.

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Neotropical Primates 21(1), July 2014 98 Figure 74. Pithecia irrorata. (a) Adult male, FMNH No. 98040, skin, Altamira, Rio Manu, Peru, and (b-c) subadult male/large juvenile, NNMN Leiden No. 409 mount and detail, Peru. Figure 75. Pithecia irrorata. (a) Captive (pet) adult male, Rio Branco, Acre, Brazil, photo by J.C. Bicca-Marques, (b) Juvenile male Pithecia irrorata pet in Iquitos, photo by Science Photo Library (http://www.sciencephoto. com/media/384772/view). a b c a b

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Neotropical Primates 21(1), July 2014 99 Figure 76. Pithecia irrorata. Adult female MNRJ No. 3317, skin and skull. a b

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100 Figure 77. Pithecia irrorata. Females: (a) adult from Ro Urubamba, Peru, photo by T. Gregory and SCBI-CCES, (b) large juvenilesubadult from Ro Urubamba, Peru, photo by T. Gregory and SCBI-CCES, and (c) a small juvenile captive, eating a cashew fruit, in the Reserva de Desenvolvimento Sustentvel Piaga-Purus, Rio Purus, Amazonas, Brazil, photo by Felipe Rossoni Cardoso. a c b

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101 a c b VANZOLINIS BALD-FACED SAKI Pithecia vanzolinii ( Hershkovitz, 1987) Synonymy 1987 Pithecia irrorata vanzolinii Hershkovitz, P. Am. J. Primatol. 12: 426; in part. Holotype. Universidade de So 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 ex ample an adult female No. 5495, skin and skull. Type locality. Brazil, Santa Cruz, Rio Eir, a southern tributary of the Rio Juru, Amazonas, for both the holotype and all paratypes. Description. Species determination is based on 36 speci mens. Males and females are similar in pelage with black dorsal fur stippled in light white to cream grizzle, with contrasting pale yellowish-bufy arms and legs. A bufy ruf matches the limbs on both males and females, but the males ruf is more distinct. A detailed description is given in Hershkovitz (1987). e descriptions by Hershkovitz and this paper are based solely on museum specimens. Males. e 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 pres ent, 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 speciesthere 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 Juru and the south bank of the Rio Tarau ac in the south-west of the states of Amazonas and Acre. ere are, however, very few documented locations for these animals, which are principally along the south-east ern side of the Rio Juru, including: Amazonas Rio Eiru, Santa Cruz, Santo Antnio, Lago Grande; Acre Serin gal 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). Table 23 Measurements for P. vanzolinii holotype and a female paratype. Measurement Male MZUSP 5491 Female MZUSP 5495 Skull (mm) Length of brain case 84.6 83.9 Width of brain case 42.1 42.3 Zygomatic arch width 57.2 57.2 Orbits outer 44.1 43.0 Nose bridge 6.8 6.3 L-orbit inner width 16.8 16.0 L-orbit inner height 17.1 16.8 Muzzle width 25.0 26.6 L-mandible length 54.3 58.8 L-mandible height 40.0 35.4 Post-crania (mm) Total length 935 740 Tail 420 465 Hind foot 120 120

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102 Specic locations. Appendix I. BRAZIL. Amazonas: Rio Jurua Lago Grande; Rio Eiru Santa Antnio, Santa Cruz. Acre : Rio Juru 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 efort should be made to gather as much information as possible to determine its range and conser vation status. Map 13. Geographic distribution of Pithecia vanzolinii. en Brazil 1. Seringal Oriente 2. Cruzeiro do Sul 3. Rio Eir, Santa Cruz (approx.) 4. Rio Eir, Santo Antnio (approx.) 5. Rio Eir, Lago Grande (approx.)

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Neotropical Primates 21(1), July 2014 103 Figure 78. Pithecia vanzolinii from Hershkovitz (1987): (a) From Figure 19 male and female face, and (b) Figure 27, original color painting by Zorica Dabich and Philip Hershkovitz, through the courtesy of Field Museum of Natural History. Figure 79. Pithecia vanzolinii. Holotype, adult male MZUSP 5491. Photos by L.K. Marsh. Figure 80. Pithecia vanzolinii. Skull of holotype MZUSP 5491. Photos by L.K. Marsh.a b

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Neotropical Primates 21(1), July 2014 104 Figure 81. Pithecia vanzolinii. Adult female paratype, MZUSP 5495.

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105 enMITTERMEIERS TAPAJS SAKIPithecia mittermeieri sp. nov. Synonymy 1987 P. i. irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425, 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 and included some of the rst eld work on sakis. He has been particularly inuential in Brazil, where he has been working since 1971 and where his eforts on behalf of Brazils amazing biodiversity have earned him the Order of the Southern Cross from the Brazilian government and the Joo Pedro Cardoso Award from the state of So 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. Hofmanns, 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 Humait (Paynter and Traylor, 1991). Paratypes: Brazil, Par, Arua, Rio Arapiunssouth of the Rio Amazonas and just west of the Rio Tapajs. BMNH No. 20.7.14.6 and MNRJ No. 3315 Rio Tapajs, Vila Braga (south of Itai tuba, left bank). INPA No. 5707 from the Municpio de Novo Aripuan, margem direita do Rio Aripuan, Lago do Cipotuba. Specimens examined. irty 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 fe males 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 dense ly covered in shorter white hairs. Wrist cufs are white un derneath and grizzled white on top. e 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. e hands and feet are white. e ruf is bright orange, but varies in density and extent among individuals. e 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 hairsvarying in intensity of whiteness depending on the ageform 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

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106 along the sides of the face and the eyebrows, and as light malar lines and lips. e extent of the light white facial hair decreases as the animals get older. e 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, ne white hairs form over the eyes extending back to a horseshoe of ne white fringe around the face. In some photos of living P. mittermeieri subadult males appear gray faced, where the white can seem obvi ous 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 ruf than males in gen eral, 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. e 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 rufs, 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 bufy to white forearms. Much older adults can be quite white in overall pelage, but unlike P. rylandsi sp. nov. males, retain their bright orange rufs. Females in the upper sections of the range south of the Amazon and between the rios Madeira and Tapajs 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 Ma deira and Tapajs, including the Rio Aripuan drainage. Current populations appear to be concentrated primarily north of Aripuan in Mato Grosso, although there is some evidence that historically they may have extended at least along the Madeira as far as the Mamor and even Rio Gua por (as per Natterer 1829 in Destacamento do Ribeiro and Rio Pacas Novos, but cf. P. rylandsi sp. nov.). e collection of a male, juvenile male, and infant in Nova Braslia, Polonoroeste, presumed to be Ji-Paran as per the gazetteers of Paynter and Traylor (1991), in 1985, demon strates some populations north of the Rio Jamar as well. Two photos of wild males taken in southern Rondnia sug gest 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 Table 24. Pithecia mittermeieri adult male holotype and male and female paratypes. Measurement BMNH 27.8.5.9.1 Male MZSP 5549 Male INPA 5707 Female MNRJ 3315 Female Skull (mm) Length of brain case 85.8 84.8 76.6 82.9 Width of brain case 43.4 44.9 42.1 47.0 Zygomatic arch width 58.4 55.7 50.0 Orbits outer 46.7 43.6 41.0 43.3 Nose bridge 8.0 7.3 6.1 8.3 L-orbit inner width 17.5 15.6 15.5 14.6 L-orbit inner height 17.0 19.0 17.4 14.4 Muzzle width 27.0 26.1 20.2 24.5 L-mandible length 57.9 56.4 50.5 51.0 L-mandible height 27.0 37.3 34.1 Post-crania (mm) Head-body 460 450 370 350 Tail 450 450 410 490 Hind foot 85 135 105 115 Ear 35 30 30 Skull articulated and not fully cleaned.

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107 the Tapajs, they extend possibly as far as Jacaracanga and perhaps further south, depending on the presence of other species (see Discussion below). Specic location. Appendix I. BRAZIL. Para : Rio Madeira Borba, Humait; Parintins Villa Bella Imperatriz, Lago Andina; Itaituba; Rio Arapiuns Arua; Rio Humait; Rio Aripuan; Rio Tapajs; Lago do Cipotuba; Villa Braga; Igarap Auara; Igarap Brabo; Igarap Amorin; Limoal; Limontuba; Santarm Fazenda Mararu. Mato Grosso: Aripuan. Rondnia: Nova Braslia. Discussion. e colloquial name for these sakis has gen erally been the Tapajs saki, and it will likely continue to be so even though the species name has been given as P. mittermeieri. e region between the rios Madeira and Tapajs has changed extensively over the last two hundred years. Railways and telegraph lines were introduced at the turn of the 19th century (instigated by Marechal Rondon). Because of highways such as the BR 364 in Rondnia com pleted 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 Rondnia 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 Comisso Rondon, or how far north of the Rio Guapor P. rylandsi sp. nov. extended orig inally, but todays distribution is likely very diferent for all of these species. So while it is possible that P. mittermeieri extended throughout the region between the rios Madeira and Tapajs to the Guapor, its exact distribution today is unknown.Figure 82. (a) Illustration of a juvenile male Pithecia mittermeieri by Graham Allen for P. Whiteld & R. Walker, and (b) Illus tration of a subadult or adult male P. mittermeieri by Alexandre Rodrigues Ferreira (1783). a b

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108 Map 14. Geographic distribution of Pithecia mittermeieri. Brazil 1. Amorin, Brabo, Limoatuba (Olalla collection, by gazetteer, Paynter Jr. & Traylor Jr. 1991) 2. Rio Arapiuns, Aru (Olalla collection, by gazetteer, Paynter Jr. & Traylor Jr. 1991) 3. Limoal near Boim (Olalla collection, by gazetteer, Paynter Jr. & Traylor Jr. 1991) 4. Parintins 5. Rio Andira 6. Lago Andira 7. Itaituba, Villa Braga 8. Igarap Auara (Olalla collection, by gazetteer, Paynter Jr. & Traylor Jr. 1991) 9. Borba 10. Rio Aripuan 11. Lago do Cipotuba 12. Humait 13. Destacamento do Ribeiro (Natterer 1883) 14. Rio Pacas Novos (Natterer 1829) 15. Nova Braslia (Polonoreste 1985) 16. Cacoal State Park 17. Chupinguala 18. Retiro do Veado Branco, Serra do Norte (Comisso Rondon) 19. Km 16 on BR 230 (approx.)

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Neotropical Primates 21(1), July 2014 109 Figure 83. Pithecia mittermeieri. Holotype BMNH No. 27.8.5.9.1, adult male, skin.

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Neotropical Primates 21(1), July 2014 110 Figure 84. Pithecia mittermeieri. Paratype INPA No. 5707, lactating subadult female, skin and skull.

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111 Figure 85 Pithecia mittermeieri. Males: (a) captive juvenile male, photo by L.C. Marigo, (b) captive juvenile male, photo L.C. Marigo, (c) captive subadult male, photo by N. Rowe, (d) captive adult male, photo by Matheus Fortunato, and captive adult male (older), photo by ZooChat.com, and (f) wild adult male in Chupinguaia, Rond nia, Brazil, by Kurazo Matheus Okada Aguilar.a e f b c d

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Neotropical Primates 21(1), July 2014 112 Figure 86. Pithecia mittermeieri. Females: (a) adult captive female, photo by L.C. Marigo, and (b) subadult female with older adult male, photo by L.C. Marigo. b a

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113 b a RYLANDS BALD-FACED SAKI Pithecia rylandsi sp. nov. 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 Whi tesides, 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; 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 Scien tist at Conservation International, Deputy Chair of the IUCN/SSC Primate Specialist Group, a member of the Brazilian Academy of Sciences, former professor of Verte brate 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 Internationals Primate Action Fund, has directly impacted primate research and conservation the world over. Dr. Rylands is a tireless aca demic 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 Ro Tahuamanu, Mucden; holotype skull AMNH No. 248723 from the Ro Nareuda. Paratypes: FMNH No. 123967 from Bolivia, Pando De partment, Porvenir, Ro 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 natu ral history museums. Description. One of the largest of the sakis. Males and fe males 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. eir dorsal coloration can be so intense with white that they take on a skunky

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114 appearance. Males in Peru appear to have almost no black showing through the heavy white hair tips. e forelimbs can be bufy 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). e hands and feet are white to of-white. e 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. is makes for the ap pearance of black stripes on the sides of the older males heads. is 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. ey lack an obvious orange or bright ruf as in P. irrorata; instead they have a black or dark brownish ruf. 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. eir forearms can be more bufy than those of the males, and this blonde-tan can extend across their chests. ey do not have distinct ruf color ation, as it is sometimes a light buf or grayish. e 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 rufs 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 Rondnia 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 Ro Madre de Dios in those areas. Specic locations with verication include: Mucden; north of the Ro Tahuamanu, but east of the Peru border; and Municpio 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 munici pality include: Bolpebra Buena Vista, Piaou, Rutina, Los Campos, El Refugio, Estacin Tahuamanu, and San Sebastian (reported not common); Filadela Pingo de Oro (rare) and north of Ro Tahuamanu, vicinity of Ro Muyumanu; Bella Flor Santa Rosa near Ro Abun, Bella Flor, Ro Manuripi, and between the ros Acre and Tahua manu; Cobija Cobija, Belo Horizonte, Tres Corazones, Puerto Rico, Tres Estrellas, Fortaleza; Porvenir Porvenir along the Ro Nareuda and Cocamita; Nuevo Esperanza Federico Roman (Compartamentos Caiman, Manoa, and Ro Negro); and Ingavi Centro Dieciocho, north of Barraca San Juan de Nuevo Mundo. Peru Mainly north of the Ro Madre de Dios, likely throughout the Reserva Territorial Madre de Dios, spe cically in the Los Amigos Conservation Concession in cluding Ro 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 Ro Madre de Dios, but this is unconrmed. eir 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.Table 25. Skull measurements for Pithecia rylandsi holotype male AMNH 248723, and paratype female KUPRI 5973. Measurements AMNH 248723 Male KUPRI 5973 Female Skull (mm) Length of brain case 63.6 77.9 Width of brain case 42.6 n/a Zygomatic arch width 54.8 51.4 Orbits-outer 42.6 40.4 Nose bridge 4.4 L-orbit inner width 15.5 L-orbit inner height 19.1 Muzzle width 21.8 L-mandible length 62.2 49.5 L-mandible height 47.9 36.4 Symphasis angle 37.0 M1-M1 mandible 21.3 M3-M3 maxillary 22.6 I-M3 maxillary 34.1 Nasal length 19.2 Nasal width 17.1 Condylobasal length 74.6 Postorbital constriction 34.1 Canine (L/R) 9.4/9.2

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115 Brazil. While it is likely that these animals occur east and north of the Rio Guapor into the states of Rondnia and Mato Grosso, and possibly as far north as the Rio Anari or west to the Rio Jamari (see Discussion, below), to be con servative in this account, their distribution will be restrict ed to the south (west) of the Rio Guapor or just north (east) along its Brazilian shores where passable by arboreal primates. Specic locations. Appendix I. BOLIVIA. Pando: Nicolas Suarez Bolpebra; Tahuamanu; Mucden; Rio Nareuda. BRAZIL. Rond nia: Cacoal Selva Park; Rio Guapor; De stacamento do Ribeiro; Rio Anari; Pedra Branca; Ritiro do Veado Branco Serra do Norte; Guajar-Mirim. PERU. Madre de Dios: Ro Madre de Dios Manu Wildlife Center; Ro 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. ese 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. is is one of the few cases in this tax onomy where there is more information on living animals than reference material. Material collected by J. Natterer in 1829 from GuajarMirim, Rio Pacas Novos (old name, now very near Gua jar-Mirim), and Rio Guapor are juveniles and dicult to place rmly in P. rylandsi; therefore, as they are thought to have been collected on the north bank of the Rio Gua por, they will be provisionally placed in P. mittermeieri (cf. P. mittermeieri). A female (MZSP No. 22895) collected in Rondnia, 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. ry landsi population in Brazil. Except for south-eastern Peru and perhaps parts of Pando in Bolivia, it is likely the population is declining through out the range. While I do not advocate more collections of these sakis, I do suggest we get as much detailed infor mation from photos or from capture-release animals as possible to determine their relationship to P. irrorata and P. mittermeieri. Map 15. Geographic distribution of Pithecia rylandsi. Peru 1. Manu Wildlife Center 2. Concesin para Conservacin, Los Amigos 3. Los Amigos Research Center Bolivia 4. L. Porter Field Site 5. Mukden 6. Ro Nareuda Brazil 7. Rio Guapor (approx. collection area both sides of river, Natterer 1829) 8. Mato Grosso do Sul (approx. collection area, Natterer 1826)

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Neotropical Primates 21(1), July 2014 116 Figure 87. Illustration of what appears to be Pithecia rylandsi drawn in 1790 by Alexandre Rodrigues Ferreira as part of his explorations into the southern Brazilian Amazon. Figure 88. Pithecia rylandsi. Holotype male, skin AMNH No. 247669 and skull AMNH No. 248723.

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117 Figure 89. Pithecia rylandsi. (a) Paratype subadult female FMNH No. 123967, skin, and (b-c) paratype skull KUPRI No. 5973. a b c

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Neotropical Primates 21(1), July 2014 118 Figure 90. Pithecia rylandsi. Adult males (a) Pando, Bolivia, photo by Leila Porter; (b) Los Amigos, Peru, photo by Bretstickypartsblog, and (c) near Manu Wildlife Center, Peru, photo by Fred Yost. a b c

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119 Figure 91. Pithecia rylandsi. Adult females (a) wild, Pando, Bolivia. Photo by Leila Porter. (bc) captive at CENP, Belm, Par. Photos by Luiz Claudio Marigo. a a b b c c

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120 Figure 92. Pithecia rylandsi, subadult female (a) captive/rehabilitation at Inti Wara Yassi, Bolivia. Photo by www.trekker.com/ shatz. com; and (b) captive, Zoo Parque Itaituba, Arlete/icker. b a

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121 enPISSINATTIS BALD-FACED SAKIPithecia pissinattii sp. nov. Synonymy 1987 Pithecia irrorata irrorata. Hershkovitz, P. Am. J. Primatol. 12: 425, in part. 2013 P. hirsuta. Van Roosmalen, M. G. M. and Van Roos malen, 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 Pri matologia 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 endan gered Brazilian primates, including the successful breeding and management of the muriqui ( Brachyteles) in captivity. Because of his genuine kindness and intellect, and his devo tion 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 Canaboua 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, Paran do Jacar, Canaboua. Hershkovitz and others assign this location to Lago do Canaboua, just south of the Rio Solimes and between the rios Purus and Madeira. Specimens examined. irty-ve 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 com pared 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. e color bands of grizzling in the dorsal pelage vary as well from whitish to bufy-cream to tan, particularly on the forearms and shoulders, often with whiter cufs at the wrist. e hind legs are often darker, brownish to blackish, sometimes with dark bands lacking the white grizzling. e ventrum is lightly covered in black hairs with a distinct ruf ranging from bright orange to dull orange/light tan, generally with lighter roots. e hands and feet are white. Males have a white crown band of bangs, hair that is dis tinct, 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. e 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 ne hair, but not nearly as much as the juvenile males east of the Rio Ma deira (P. mittermeieri ). Females. Females often have more grizzling in the dorsal pelage than malesthe 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. e b

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122 ruf is present with dark brown base hairs ending in lighter tan to bufy bands. ey often have darker wrist cuf bands above the hands. e faces of adult females are covered in white soft hairs that cover the forehead, cheeks, and eye brows. 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. e white malar lines are more obvious and often shaggier than in males. e skin above and below the eyes is light pink ish, 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 tour ists on the Rio Juma (at Juma Jungle Lodge) as recently as 2009 (Figure 96, 98 a,b). Pithecia pissinattii are most simi lar 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 rufs. However, ju venile and subadult male P. mittermeieri are very diferent from those of P. pissinattii. e 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, zygo matic arch width 55.1, outer span of orbits 42.5, nasal con striction 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 Solimes in the northern area between the rios Purus and Madeira, including Autazes, Lago do Canaboua, 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. Specic locations. Appendix I. BRAZIL. Amazonas: So limes Parana do Jacar, Canaboua, Lago Canaboua; Rio Purus Lago Jari, Itaboca; Autazes; Rio Juma Juma Lodge; Rio Madeira Rosarinho, Lago Miguel, Lago Sam paio; Manaquiri. Discussion. Pithecia pissinattii sakis may occur east of the Rio Madeira, and gene ow 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 spe cies, particularly in their lower, wider reaches.Table 26. Reported measurements for P. pissinattii from Paran do Jacar, Canaboua, Brazil. Po s tcr ania (c m) He a d body T a il Hind foot E a r Males BMNH 25.12.11.2 5548 BMNH 26.5.5.7 54 48 BER 33937 51 52 BER 91314 (SAM) 50 47 BMNH 27.8.11.19 (Juv) 4542 MNRJ 3313 45 48 11.5 2.4 MNRJ 3316 50 48 12.5 2.3 SEN 6917 47 47 NHRM A64-0097 49 48 Females NHRM 3314 50 45 11.8 2.4 BMNH 26.5.5.8 51 49 BMNH 27.8.11.14 5049 BMNH 25.12.11.4 4345 BMNH 27.8.11.13 5347 BMNH 27.8.11.16 5148 BMNH 25.12.11.3 5248 BMNH 27.8.11.20 4143

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123 Map 16. Geographic distribution of Pithecia pissinattii. Brazil 1. Itaboca 2. Rio Jari 3. Canaboua (by gazetteer Paynter Jr & Traylor Jr. 1991) 4. Manaquiri 5. Juma Lodge 6. Autazes 7. Rosarinho 8. Lago Sampaio 9. Lago Miguel

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Neotropical Primates 21(1), July 2014 124 Figure 93. Pithecia pissinattii. Holotype BMNH No. 26.5.5.7, adult male, skin.

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125 Figure 94. Pithecia pissinattii. Holotype BMNH No. 26.5.5.7, adult male, skull.

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Neotropical Primates 21(1), July 2014 126 Figure 95. Pithecia pissinattii. Paratype BMNH No. 27.8.11.13, adult female skin and skull.

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Neotropical Primates 21(1), July 2014 127 Figure 96. Adult male Pithecia pissinattii at Juma Jungle Lodge, Rio Juma, Brazil. Photo by Crijnfotin via Flickr. Figure 97. Pithecia pissinattii subadult male from Manaquiri. Photo by Ricardo Sampaio.

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128 Figure 98 Pithecia pissinattii. (a) Adult female, photo by Trip Wow Website and (b) juvenile female, photo by Crijnfotin via Flickr, both at the Juma Jungle Lodge, Rio Juma, Brazil. a b

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129 b I think it is important to emphasize my goals with this monograph, as well as the goals of taxonomy in general. Groves (2001) said: Taxonomy, like other elds of biology (ecology, ethology, physiology, genetics), is a dynamic sci ence. Classications are not engraved in stone, nor should they be; it is unfortunate that advances in the taxonomic eld, unlike those in ecology and other disciplines, often require changing the names we give to species [] but that is the way it must be, and the irritation felt [] will pass quickly. Indeed, new predictions, to be tested in the eld, may well emerge from the reclassication (p.5). It is exactly my intent to create a hypothesis for the taxon omy of Pithecia so that we can continue to piece together the story, historic and present-day in the eld, to give us greater detail for understanding this fascinating group of monkeys. Ultimately, the goal is to conserve as many saki species in the wild as possible, and that is only achievable with a working taxonomy. My rationale for this work was to tame the madness of Pithecia taxonomy, not to perfectly corroborate every specimen or living saki out there. I think this treatment comes closeeven though in many cases I was a lumping splitter, I feel I gave as much rigor as possible to the determinations with the data at hand.en Taxonomic hygiene I mentioned in the Introduction the trouble with com parative studies, especially in genetics, that are lax in as suming that P. monachus, P. irrorata, or other previous catch-all species are well-dened and valid taxonomic units. Even cladistic relationships between genera, such as those comparing Pithecia to other pitheciids, one would think to be safe. However, if samples, specimens or individuals are used from more than one geographic location, the lack of a rigorous taxonomy makes it entirely possible that more than one species is being lumped together to represent the Pithecia unit. Researchers in cladistics, genetics, and morphology rarely report exactly what specimens were used, where they were collected, or on what grounds they are determining the species name. ose that do report detailed information on their test subjects, even if the taxonomy changes are at least able to clearly compare their work with others in the future. Bortolus (2008) found that biology, ecology, biodiversity, genetics, and like disciplines, tend to disregard reporting the taxonomic pedigree. is occasions a cascade of errors with negative consequences for the development of scientic knowledge, as well as for biodiversity and human welfare. He concluded that, a single incorrect taxonomic identica tion has a great potential to be assimilated into many difer ent biological and ecological studies and then in several envi ronmental management studies and programs, multiplying its impact synergistically. He maintained that these kinds of errors are likely to have a variety of negative consequenc es, such as identifying a population as homogeneous and monospecic when it should actually be recognized as a complex assemblage with distinct geographic distributions. At a minimum, cascading errors in taxonomy in our eld are annoying and inaccurate, at worse they are afecting our ability to do good science, conserve species, or get funding. Bortolus correctly argues that there is a real need for reli able taxonomic conrmation of the diferent taxa we work with through updated methods and perspectives. He main tains, Ecologists usually have the feeling that taxonomic assistance may not be needed when working in areas that have been studied for decades (or centuries) by many other ecologists. However, it is clear that just because many other ecologists have been working in the exact same place we are now does not guarantee that the existing species assemblag es are currently the same or prevent cryptic non-native species or genotypes from being introduced and unnoticed. It is clear that we can easily substitute primatologists for his ecologists as we all sufer the same biases. And while we may not necessarily have cryptic species, we certainly might have more than one species within an historically named taxa, or, as is the case with Pithecia, species whose juvenile members have such vast color variations that in the past they were called out as new species. It is clear in primatology, as it is in biology generally, that taxonomy is all too often seen as a dead science. I myself as a trained eld ecologist could not think of anything worse than having to ribe through dead animals in the dusty stacks of a museum. But the lesson learned was a valuable one: we are making more of a mess by perpetuat ing mistakes than consulting our taxonomists when we do our research. Bortolus says this: A main causal factor in the error cascade is that even though we recognize taxonomy as an important part of biology as a whole, during the last decades taxonomists have lost consideration from the rest of the scientists. If ecologists disregard or underestimate the work of taxonomists, then they must show reliability and excellence in their taxonomic identications, and this is not what presently seems to be occurring.en The path forward So what can we do? I concur with Bortolus that in an ideal world we would be sure to consult with taxonomists directly and involve or consult them about the diferent taxa in our studies before we publish so we know exactly what species we are studying, comparing, or reporting on. However, in our eld I can literally count on less than one hand the number of trained taxonomists at our disposal. Which means we all need to practice taxonomic hygiene when we publish our work, so that at a minimum we can accurately compare our ndings. D

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130 To do this I suggest the following for all studies of primates. 1. Add a voucher photograph. is is especially impor tant if you are studying Pithecia, but is needed for any species. If your document is published online, make sure it is in color. If your specimen is deadeither as a skin, skeletal material, uid, tissue, DNA, mount, or otheradd a photo of the living source, if possible. If it is not possible to obtain a photo, then a detailed description of the once-living animal could suce. 2. Provide a specic location for where you are working or where your (living/dead) subject is from. Field pri matologists tend to be good at this since they need to let us know where their eld sites are. But an extra level of detail would be nice when speaking about study animals even in situ, such as sides of a river (left or right are old conventions, use ordinates), or how far from a river you traveled and in what direction you were when you (encountered, heard, studied, took samples from) the species. 3. Report as much information as you have on samples. If doing genetics, cladistics, or other experiments: specimen numbers, collection dates, locations, col lectors, museums of origin, where you got the tissue sampleanything you can report is very important. If you receive samples from a catalog or bio supply house, it may take some doing, but track down the necessary source information. If you believe you have a new species of primate, there are some immediate things to do before dashing of a publica tion about it. 1. Contact a taxonomist. ere are many subject matter experts, but contacting one of our known primate tax onomists and have them direct you is so important. And if you do not know a taxonomist, look one up: every natural history museum has one. 2. Do not kill anything (not at rst anyway). Get good photos, maybe dart it and take material for analysis, but do not take a voucher until it is clear you have a new species; 3. Do the work. If no one has done a paper (like this one) recently, you might need to track down all of the museum specimens, living photos, eld sites, maybe do more eld workbefore you can rightfully declare a new species. In conclusion, I implore all of us to do a better job in the nomenclature and taxonomy of Primates. Because it cannot be said enough, a reliable taxonomy is among the major factors guaranteeing the accurate identication of biodiversity hotspots, wilderness areas, endangered species, and areas of greatest concern that impact all of the animals we care so much aboutand besides, researchers 200 years from now will thank you. Summary In this revision, there are ve original species, three species elevated from subspecic rank, three historic species rein stated, and ve newly described species. e total number of Pithecia species is 16. Map 17 illustrates the full range of Pithecia in the Amazon Basin and Shield countries. Figure 99 is a rough distribu tional schematic, and Figures 100 and 101 provide side-byside representation of all species and schematics of all spe cies described in this publication. Appendix III provides an interpretation based on this taxonomy of the Hershkovitz (1987) paintings by Zorica Dabich, which recently became available in color for our use through the courtesy of Field Museum of Natural History.

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131 Map 17. e approximate distributions of the saki monkeys, Pithecia, following the classication proposed. Map by Stephen D. Nash.

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132 Figure 99. Schematic of all Pithecia species. Illustration by Stephen D. Nash.

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133 Figure 99. Schematic of all Pithecia species. Illustration by Stephen D. Nash. Figure 100. Male and female faces of all Pithecia species in a roughly geographic arrangement. Illustration by Stephen D. Nash.

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134 Figure 101. e species of Pithecia, male and female, full bodies, in roughly geographic arrangement. Illustration by Stephen D. Nash.

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135 Acknowledgements A project like this cannot happen without a lot of support, help, and condence from my colleagues. So, while I will start with a blanket statement about my appreciation for everyone literally around the world who helped meand truly I cannot thank each of you enoughI will also do my best below to call out specic persons and institutions. First, I absolutely must make a fuss over Colin Groves, Anthony Rylands, and Russell Mittermeier for not only schooling me in taxonomy, but for sharpening my skills as a scientist, for being clear, unwavering guidance through this maze, and for being good friends. So much gratitude and love to you guys! ank you over and over to Stephen Nash for all of his patience in illustrating everything in this volume. It was such a privilege to work and to share comic book fandom with you! None of this work would have been possible without the gracious support of Conservation International, Primate Action Fund, the Margot Marsh Biodiversity Founda tion, Primate Conservation Inc. and Noel Rowe especially, St. Louis Rainforest Advocates, and Global Conservation Institute. In particular, I want to thank Jeanne Fair for all of her help and support over the years with this project. ank you, thank you, thank you! I am also grateful for helpful comments over the years from Marilyn Norconk, omas Deer, Andrew Kitchener, Jos de Sousa e SilvaJnior Cazuza, Mrio de Vivo, Rolando Aquino, Luis Albuja, and Erwin Palaciosmuchsimas gracias, muito obrigada, and thank you all again so very much. Austria. Vienna. Much thanks to Simon Englberger and Alice Schuhmacher for their excellent work and photos from the Naturhistorisches Museum Wien. Bolivia. Many thanks to Leila Porter and Rob Wallace for all of the helpful information, and photos of animals from the eld. Brazil. Belm: Museu Paraense Emlio Goeldi staf and stu dents (including Paulo Henrique), but particularly huge thanks to the late Liza Viega for helping me and letting me stay in her lovely apartment, and Cazuza (Jos de Sousa e SilvaJnior) for everything; and the Centro Nacional de Primatas to the staf and helpful curators. Manaus: Instituto Nacional de Pesquisas da Amaznia, especially Manoela Borges, Fabio Rhe, and Wilson Spironello, and the staf of the Projeto Dinmica Biolgica de Fragmentos Florestais (PDBFF) Ze (Jos Luis Camargo) and especially Bryan Lenz (thanks for the hammock!). Rio de Janeiro: Museu Nacional Rio de Janiero, with a full heart and world of gratitude to Joo Alves Oliveira and Leandro Salles; Jardim Zoolgico do Cidade do Rio de Janeiros Curator of Mammals and primate keepers; Centro de Primatologia do Rio de Janeiro, with special thanks to Alcides Pissinattiyou are truly one of our greats. So Paulo: Museu de Zoologia da Universidade de So Paulo, many, many thanks to Mrio de Vivo and all of his wonderful staf and students; So Paulo Zoo thanks to Maria Cristina Marques for the behind the scenes tour. Colombia. Personal thanks for incredible help to Xyo mara Carretero-Pinzn for work in the museums. Bogot: ank you to the curators and directors in the collections of: Instituto de Investigacin de Recursos Biolgicos Alexander von Humboldt, Universidad Nacional de Colombia Instituto de Ciencias Naturales, Museo de Historia Natural, Universidad de los Andes, and the Coleccin Manuel Ruiz-Garca de la Ponticia Universidad Javeriana. Ecuador. Quito: All of the staf and students for the TBS Oce at Universidad San Francisco de Quito, but particu larly Kelly Swing and David and Consuelo Romo; the col lection at Ponticia Universidad Catlica del Ecuador; spe cial thanks to Luis Albuja for allowing us access to the col lection at Escuela Politcnica Nacional; Nataly Orbe Ortiz at Museo Amaznico. Napo: special thanks to Franklin, Jendry, and Romero and their families who worked with me and my eld staf: Shayna Whitaker, Cindy orstad, and Domingo Canales Espinosa to Yarina, Yuturi, and Comuna Pamiwa Kocha; Tiputini Biodiversity Station: huge thanks to Bette Loiselle for having me work there ini tially back in 2001, thanks to Tony DiFiore for early help, and to Jose Hidalgo for eld and museum work. Pastaza: thanks to Jarol Fernando Vaca for letting us be some of the rst visitors at Shiripuno Lodge. Tena: thank you for the personalized visit at AmaZOOnico. France. Many thanks for gracious help from Anne Previato, Geraldine Veron and Jacques Cuisin of Museum dHistoire naturelle de Paris. e holotype vault is so impressive! Germany. Berlin: Peter Giere and Frieder Mayer, thank you so much for your help in the collections of the Museum fur Naturkunde. I really enjoyed your museum. Bonn: Many thanks to Gustav Peters for interesting discussions and data from Zoologisches Forschungsmuseum Alexander Koenig and the Cologne Zoo. Frankfurt: Grateful thanks to Katrin Krohman at Naturmuseum Senckenberg. You made me feel so welcomed. Munich: Richard Kraft, I cannot thank you enough for allowing me to stay and study in the Zoologische Staatssammlung. Your expertise and friendship was greatly appreciated. Hungary. ank you to Gabor Csorba for sending photos from the collection in the Hungarian Natural History Museum. Italy. I appreciated so much meeting Dr. Cecilia Veracini in Torino, and getting photos and data from the collection in the Museo di Storia Naturale di Firenze (Florence) from her. Japan. Many thanks for your hard work to Hikaru Waka mori and to Masanasu Takai for photos and data from the Kyoto Primate Research Institute. Nederlands. Many thanks to Madame Bik for her hos pitality for my stay in her home. Huge thanks to Chris Smeenk for his help and guidance and to Hein von Grouw for allowing me access to the collection at the Nationaal Natuurhistorisch Museum, Leiden Naturalis.

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136 Peru Galaxies of thanks to Fanny Cornejo for all of your help and friendship over the years, but in particular for our tour through Peru! Also a great many thanks to Fanny Fernandez Melo for letting me stay in your home. Iquitos: Huge thanks to Rolando Aquino for letting me study your specimens at the Proyecto Peruano de Primatologa de la Universidad Nacional Manuel Moro Sommo. I look forward to one day working with you! ank you to the staf of the Museo de Zoologa FCB-UNAP for allowing us access to all of your skulls, and special thanks to Csar Terrones Ruiz and Fanny Cornejo for help measuring them; thank you to all of the staf at Isla de los Monos and Pilpintuwasi for allowing us access to your animals. Lima: Hundreds of thanks to the Mastozoologa Peruana group, Sandra Mori, Carlos Tello, all of the students and staf, and particularly director Victor Pacheco of the Museo de Historia Natural Javier Prado de la Universidad Nacional Mayor de San Marcos for allowing me to work in your collection; many grateful thanks to the curators and keepers of Parque de las Leyendas and Parque Zoolgico de Huachipa for your help and behind the scenes tours. Sweden. I am so grateful for the help and friendship of Olavi Gronwall and the staf of the Naturhistoriska Riks museet, Stockholm. I loved all of the mandatory staf breaks and lunches! Switzerland. I really enjoyed working at the University of Zrich-Irchel Anthropological Institute and Museum. Lots of thanks to all of the staf and students who helped me while I was there. United Kingdom. London: Great thanks to Paula Jenkins and the mammal division of the British Museum of Natural History. Such an amazing collection! Also thanks to Malcom Fitzpatrick of the London Zoo for the personal ized tour. Scotland: Great thanks to Andrew Kitchener for your kind guidance and help at the Royal Scottish Museum in Edinburgh and at the Edinburgh Zoo. I look forward to working with you again! United States. Berkeley, CA: Special thanks to Francisco Boavista Pontual (Chico) for being my eyes while work ing at the Berkeley Natural History Museum, University of California. Chicago, IL: ank you to my long time friends and colleagues Robin Foster and Tyana Watcher for ini tially steering me to the right people, namely Bill Stanley and Michi Schulenberg of e Field Museum for my mul tiple visits. It was so amazing to work in the collection that Philip Hershkovitz once curated, and to see original saki paintings from his publications. Los Angeles, CA: anks to Jim Dines at the Los Angeles County Museum of Natu ral History for being my rst museum on the long path to saki discovery! New York, NY: ank you for having me several times especially to Eileen Westwig of the American Museum of Natural History. Special thanks to Brandon Pearson for all of your work in the collection to ll in some of the data that crashed. Washington, DC: ank you so much to Linda Gordon and David Schmidt of Smithson ian Institution. And nally, to many, many individuals the world over who ofered advice, photos, data, help, and support, includ ing (but not limited) to: Azra Marghoob, Mark Bowler, Eckhard Heymann, Anne Savage, Susan Palminteri, Leila Porter, Tremie Gregory, Jan Vermeer, Ricardo Sampaio, Horcio Schneider, Jean Boubli, Andrew Young, Andrew Johns, Jlio Csar Bicca-Marques, Felipe Ennis, Eduardo Paschoalini, Stephen Ferarri, Javier Garcia, Janice Chism, Alexey Tishechkin, Bruce Junek, Dave Freeman, Shawn Heick, Joanna Buerling, Michael Turco, and the Pitheci ine Action Group.

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137 Br Rr Adams, D. B. and Erhart, E. M. 2009. Diferential response to predators in the Grays bald-faced saki monkey ( Pithecia irrorata): a playback experiment. Am. J. Phys. Anthropol. Suppl. 48: 75. Allen, J. A. 1914. New South American monkeys. Bull. Am. Mus. Nat. Hist. 33: 647-655 Allen, J. A. 1916. List of mammals collected for the Ameri can Museum in Ecuador by William B. Richardson, 1912. Bull. Am. Mus. Nat. Hist. 35, Article XIII. p. 125: #69. Alverson, W., Moskovits, D. and Shopland, J. 2000. Rapid Biological Inventories, Bolivia: Pando, Ro Tahuamanu. e Field Museum, Chicago, IL, pp.47. Aquino, R., Alvarez, J. and Mulanovich, A. 2005. Diver sity and conservation status of primates in the Sierras de Contamana, Peruvian Amazonia. Rev. Peru. Biol. 12: 427. Aquino, R., Cornejo, F. M., Lozano, E. P. and Heymann E. W. 2009. Geographic distribution and demography of Pithecia aequatorialis (Pitheciidae) in Peruvian Amazonia. Am. J. Primatol. 71: 964. Aquino R. and Encarnacin, F. 1994. Primates of Peru/Los Primates del Per. Primate Report (40): 1. Ayers, J. M. and Milton, K. 1981. Survey of primates and their habitat on the Tapajs River, Brazil. Bol. Mus. Paraense Emilio Goeldi Nova S rie Zoologia 11: 3. Ayres, J. M. and Clutton-Brock, T. H. 1992. River bound aries and species range size in Amazonian primates. Am. Nat. 140: 531. Auricchio, P. 2000. Phylogeny of saki monkeys, Pithecia. Neotrop. Primates 8(1): 51. Auricchio, P. 2010. A morphological analysis of some spe cies of Callicebus omas, 1903 (Pitheciidae Callicebi nae). Neotrop. Primates 17(2): 47. Barnett, A., Borges, S. H., de Castilho, C.V., Neri, F. M. and Shapley, R. L. 2002. Primates of the Ja National Park, Amazonas, Brazil. Neotrop. Primates 10: 65. Bartecki, U. and Heymann, E.W., 1987. Uber Schweifafen in Peru. Zeit. Kolner Zoo 30: 79. Bates, H. W. 1892. e Naturalist on the River Amazons John Murray, London. 336pp. Bennett, C. L., Leonard, S. and Carter, S. 2001. Abun dance, diversity, and patterns of distribution of primates on the Tapiche River in Amazonian Peru. Am. J. Primatol. 54: 119. Bodini, R. and Prez-Hernndez, R. 1995. Distribution of the species and subspecies of cebids in Venezuela. Fieldiana Zool., n.s. 39: 231. Bodmer, R. E., Eisenberg, J. F. and Redford, K. H. 1997. Hunting and the likelihood of extinction of Amazonian mammals. Cons. Bio. 11: 460. Bortolus, A. 2008. Error cascades in the biological sciences: the unwanted consequences of using bad taxonomy in ecology. Ambio 37(2): 114. Branch, L. C. 1983. Seasonal and habitat diferences in the abundance of primates in the Amazon (Tapajs) National Park, Brazil. Primates 24: 424. Brooks, T. and Helgen, K.M. 2010. A standard for species. Nature, Lond. 467: 540. Burmloop, A., Homburg, I., Peetz, A. and Riehl, R. 1994. Gular scent glands in adult female white-faced saki, Pithecia pithecia and eld observation on scent marking behavior. Folia Primatol. 63: 212. Buzs1, B. and Csorba, G. 2010. Catalogue of recent pri mates in the Hungarian Natural History Museum. Ann. Mus. Natl. Hungar. 20: 229. Buchanan, D. B., Mittermeier, R. A. and Van Roosmalen, M. G. M. 1981. e saki monkeys, genus Pithecia. In Ecology and Behavior of Neotropical Primates A. F. Coimbra-Filho and R. A. Mittermeier (eds.), pp.391 417. Academia Brasileira de Cincias, Rio de Janeiro. Bufon, M. 1789. Histoire Naturelle Generale et Particuliere, A LHistoire des Animaux Quadrupedes, Supplement Tome Septieme. LImprimerie Royale: Paris; pp. 113. Burton, F. 1995. e Multimedia Guide to the Non-Human Primates. Prentice Hall, NJ. Collar, N. J. 1997. Taxonomy and conservation: chicken and egg. Bull. B. O. C. 117: 122. Cunningham, E. P. and Janson, C. H. 2006. Pithecia pithecias behavioral response to decreasing fruit abundance. Am. J. Primatol. 68: 491. Cunningham, E. and Janson, C. 2007. Integrating information about location and value of resources by whitefaced saki monkeys ( Pithecia pithecia). Anim. Cogn. 10: 293. da Rocha, R., Harada, M. and Schneider, H. 1992. ABO Blood groups in Cebidae (Platyrrhini, Primates) from the Rio Jamari region. Primates 33: 391. de Oliveira Azevedo Lopes, M. A. and Rehg, J.A. 2003. Observations of Callimico goeldii with Saguinus imperator in the Serra do Divisor National Park, Acre, Brazil. Neotrop. Primates 11: 181. de la Torre, S. 2000. Primates de la Amazonia del Ecuador Simbioe, : Quito. de oisy, B., Renoux F. and Julliot, C. 2005. Hunting in northern French Guiana and its impact on primate com munities. Oryx 39: 149. Deer, T. R. 2004. Primates of Colombia. Conservation International, Washington DC. Detwiler, K. M. 2004. Hybridization between red-tailed monkeys (Cercopithecus ascanius) and blue monkeys ( C. mitis) in East African forests. In: M. E. Glenn, and M. Cords (eds.), e Guenons: Diversity and Adapta tion in African Monkeys pp.79. Kluwer Academic / Plenum Publishers, New York.

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Seed predation by monkeys and macaws in eastern Ven ezuela: preliminary ndings. Primates 38: 177. Nowak, R. M. 1997. Sakis. In: Walkers Mammals of the World. Website: . Olalla, A. M. 1935. El genero Sciurillus representado en la amazonia y algunas observaciones sobre el mismo. Rev. Mus. Paulista, S o Paulo 19: 425. Olalla, A. M. 1937. I. Um viaje a pesquizas zoologicas hacia el Rio Jurua [sic], Estado del Amazonas, Brasil. 1936. II. Notas de campo: observaciones biolgicas. Rev. Mus. Paulista, So Paulo 23: 235. Oliveira, J. M. S., Lima, M. G., Bonvincino, C., Ayres, J. M. and Fleagle, J. G. 1985. Preliminary notes on the ecology and behavior of the Guianan saki ( Pithecia pithecia Linnaeus 1766; Cebidae, Primata). Acta Amazonica 15: 249. Pacheco, J. F. 1995. New distributional records for some birds from vrzea forest at Mamirau Reserve, western Brazilian Amazonia. Ararajuba 3: 83. Palminteri, S. and Peres, C. A. 2012. Habitat selection and use of space by bald-faced sakis ( Pithecia irrorata) in southwestern Amazonia: lessons from a multiyear, multi group study. Int. J. Primatol. 33: 401. Patterson, B. D. 1991. Mammals in the Royal Natural His tory Museum, Stockholm, collected in Brazil and Bolivia by A. M. Olalla during 1934. Fieldiana Zool., n. s. (66): 1. Patton, J. L., da Silva, M. N. F. and Malcolm, J.R. 2000. Mammals of the Rio Juru and the evolutionary and eco logical diversication of Amazonia. Bull. Am. Mus. Nat. Hist. 244: 1. Paynter, R. A. and Traylor Jr., M. A. 1977. Ornithological Gazetteer of Ecuador. President and Fellows of Harvard College, Boston, MA. Paynter, R.A., Traylor Jr., M. A. 1981. Ornithological Gaz etteer of Colombia President and Fellows of Harvard Col lege, Boston, MA. Paynter, R.A. 1982. Ornithological Gazetteer of Venezuela. President and Fellows of Harvard College, Boston, MA. Paynter, R.A. and Traylor Jr., M.A. 1991. Ornithological Gazetteer of Brazil {2 vols.). President and Fellows of Harvard College: Boston, MA. Paynter, R. A., Traylor Jr., M. A., and Winter, B. 1975. Ornithological Gazetteer of Bolivia. President and Fellows of Harvard College, Boston, MA. Pelzeln, A. von, 1883. Brasilische Saugethiere. Resultate von Johann Natterers Reisen in den Jahren 1817 bis 1835. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 33(Beiheft): 1/13 [ Pithecia]. Pereira, M. E. and Fairbanks, L. A. 1993. Juvenile Primates: Life History, Development, and Behavior. Oxford Univer sity Press, Oxford. Peres, C. A. 1993. Notes on the ecology of bufy saki mon keys (Pithecia albicans, Gray 1860): a canopy seed-preda tor. Am. J. Primatol. 31: 129. Peres, C. A. 1997. Primate community structure at twenty western Amazonian ooded and unooded forests J. Trop. Ecol. 13: 381. Pitman, N., Loyola, M., Salas, K., Vigo, G. and Lutz, D. 2005. Bibliografa Verde de Madre de Dios, Version 4.1. 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142 Rylands, A. B. and Mittermeier, R. A. 2009. e diver sity of the New World primates (Platyrrhini). In: South American Primates: Comparative Perspectives in the Study of Behavior, Ecology, and Conservation In: P. A. Garber, A. Estrada, J. C. Bicca-Marques, E. W. Heymann and K. B. Strier (eds.), pp.23. Springer, New York. Rylands, A. B., Mittermeier, R. A. and Rodrguez-Luna, E. 1995. A species list for the New World Primates (Platyr rhini): distribution by country, endemism, and conserva tion status according to the Mace-Lande system. Neotrop. Primates 3(suppl.):113. Rylands, A. B., Schneider, H., Langguth, A., Mittermeier, R. A., Groves, C. P. and Rodrguez-Luna, E. 2000. An assessment of the diversity of New World primates. Neo trop. Primates 8: 61. Rylands, A. B., Matauschek, C., Aquino, R., Encarnacin, F., Heymann, E. W., de la Torre, S. and Mittermeier, R. 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M., Encarnacin, F., Montoya, E. and Salzano, F.M. 1995. Biochemical diversity and genetic distances in the Pitheciinae subfamily (Primates, Platyrrhini). Primates 36:129. Schreber, J. C. D. 1775. Die Sugethiere, in Abbildungen nach der Natur mit Beschreibungen, Erster eil. Der Mensch. Der Afe. Der Maki. Die Fledermaus. 190pp. [Tab. XXXII. Simia Pithecia LINN. BUFF. XV.] Erlan gen. (Walther). Sclater, P. L. 1879. e Vertebrated Animals Now or Lately Living in the Gardens of the Zoological Society of London 7th edition, pp. 30. Zoological Society London, London. Setz, E. Z. F., Enzweiler, J., Solferini, V. N., Amndola, M. P. and Berton, R. S. 1999. Geophagy in the golden-faced saki monkey (Pithecia pithecia chrysocephala) in the Central Amazon. J. Zool., Lond. 247: 91. Setz, E. Z. F. and Gaspar, D. de A. 1997. Scent-marking behaviour in free-ranging golden-faced saki monkeys, Pithecia pithecia chrysocephala: sex diferences and context. J. Zool., Lond., 241: 603. Silva-Jnior, J. S. and Figueiredo, W. B. 2006. Pitheciine taxonomy and geographic distribution. Paper presented at XXI Congress of the International Primatological So ciety, Entebbe, Uganda. Smeenk, C. 2009. Has one of Captain Cooks possums landed in Leiden? e possible holotype of Pseudocheirus peregrinus (Boddaert, 1785). Zool. Med. Leiden 83: 723. Soini, P. 1972. e capture and commerce of live mon keys in the Amazonian region of Peru. Int. Zoo Yearb. 12: 26. Soini, P. 1986. A synecological study of a primate commu nity in Pacaya-Samiria National Reserve, Peru. Primate Conserv. (7): 63. Spix, J. B., von. 1823. Simarium et Vespertiliarum Brasilienses Species Novae. F.S. Hbschmann, Munich. pp. 14, pls. 9. Stephens, L. and Traylor Jr., M. A. 1983. Ornithological Gazetteer of Peru. President and Fellows of Harvard Col lege, Boston, MA. Stephens, L. and Traylor Jr., M. A. 1985. Ornithological Gazetteer of the Guianas. President and Fellows of Har vard College, Boston, MA. Tappen, N. C. 1969. Primate skull collections in some Eu ropean museums. Curr. Anthropol. 10: 62. Tate, G. H. H. 1939. Pithecia Desmarest. Mammals of the Guiana Region. Bull. Am. Mus. Nat. Hist. 76(5):220. omas, O. 1903. Notes on South-American monkeys, bats, carnivores, and rodents, with descriptions of new species. Ann. Mag. Nat. Hist., 7th series 12: 455. omas, O. 1927a. e Godman-omas Expedition to Peru. V. On mammals collected by Mr. R. W. Hendee in the Province of San Martn, N. Peru, mostly at Yurac Yacu. Ann. Mag. Nat. Hist., 9th series 19: 361. omas, O. 1927b. e Godman-omas Expedition to Peru. VI. On mammals from the Upper Huallaga and neighbouring highlands. Ann. Mag. Nat. Hist., 9th series 20: 594. omas, O. 1928. e Godman-omas expedition to Peru .VII. e mammals of the Rio Ucayali. Ann. Mag. Nat. Hist., 10th series 2: 249. ompson, C. L. and Norconk, M. A. 2011. Withingroup social bonds in white-faced saki monkeys ( Pithecia pithecia) display male-female pair preference. Am. J. Primatol. 73: 1051. ompson, C. L. Whitten, P. L. and Norconk, M. A. 2011. Can male white-faced saki monkeys ( Pithecia pithecia) detect female reproductive state? Behaviour 148: 1313. Timm, R. M., Ramy II, R. R., and the Nomenclature Committee of the American Society of Mammalogists. 2005. What constitutes a proper description? Science 309: 2163. Tirira, S. D. (ed.). 2001. Libro Rojo de Los Mamferos de Ecuador. Sociedad para la Invetigacin y Monitoreo de la Biodiversidad Ecuatorania (SIMBIOE) / Ecociencias / Ministerio del Ambiente / UICN. Serie Libros Rojos

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143 del Ecuador, Tomo 1. Publicacin Especial sobre los Mamferos del Ecuador. 236pp. Ulloa, R. 1988. Estudio Sinecolgico de Primates en la Reserva de Produccin Faunstica Cuyabeno, Amazona Ecuatoriana. Tesis de licenciatura, Ponticia Universidad Catlica del Ecuador, Quito. Van Roosmalen, M. G. M., van Roosmalen, T. and Mit termeier, R. A. 2002. A taxonomic review of the titi monkeys, genus Callicebus omas, 1903, with the de scription of two new species, Callicebus bernhardi and Callicebus stephennashi, from Brazilian Amazonia. Neo trop. Primates 10(Suppl.): 1. Vanzolini, P. E. 1993. As viagens de Johann Natterer no Brasil, 1817. Pap. Avuls. Zool., So Paulo 38: 17. Vi, J.-C., Richard-Hansen, C. and Fournier-Chambrillon, C. 2001. Abundance, use of space, and activity patterns of white-faced sakis ( Pithecia pithecia) in French Guiana Am. J. Primatol. 55: 203. Veiga, L. M., Mittermeier R. A. & Marsh, L. K. 2008. Pithecia albicans. In: IUCN 2013. IUCN Red List of reatened Species. Version 2013.2. . Downloaded on 29 April 2014. Veiga, L. M. & Marsh, L. K. 2008a. Pithecia irrorata In: IUCN 2013. IUCN Red List of reatened Species. Ver sion 2013.2. . Downloaded on 29 April 2014. Veiga, L. M. & Marsh, L. K. 2008b. Pithecia pithecia. In: IUCN 2013. IUCN Red List of reatened Species. Ver sion 2013.2. . Downloaded on 29 April 2014. Voss, R .S. and Fleck, D. W. 2011. Mammalian diversity and Matses ethnomammology in Amazonian Peru. Part I: Primates. Bull. Am. Mus. Nat. Hist. (351): 1. Wgele, H., Klussmann-Kolb, A., Kuhlmann, M., Haszprunar, G., Lindberg, D., Koch, A., Wgele, J. W. 2011. e taxonomist an endangered race. A practical pro posal for its survival. Frontiers Zool. 8: 25. Wagner, A. 1847. Mammalis, Simiae. In: Reports on Zoology for 1843, 1844, translated from the German by Busk, G., Tulk, A., Haliday, A. H. e Ray Society, London, p. 20. Wagner, A. 1848. Beitrage zur Kenntniss der Saugethiere Amerikas. Dritte Abtheilung. Vierte Ordnung. Aen. Ab handlungen der Mathematisch-Physikalischen Classe der Koniglich Bayerischen Akademie der Wissenschaften 5: 405. 1 pl. Wakeham-Dawson, A., Morris, S. and Tubbs, P. 2002. Type specimens: dead or alive? Bull. Zool. Nomenclat. 59:282. Walker, S. E. 2005. Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela. Am. J. Primatol. 66: 369. Wallace, A. R. 1889. A Narrative of Travels on the Amazon and Rio Negro with an Account of the Native Tribes, and Observations on the Climate, Geology, and Natural history of the Amazon Valley. 2nd edition. Ward, Lock and Co., London. pp. 311, 328. Wallace, A. R. 1900. e hairy saki. Studies Scientic and Social 1: 172. Macmillan & Co., London. Wiley, R. H. 2010. Alfonso Olalla and his family: the or nithological exploration of Amazonian Peru. Bull. Am. Mus. Nat. Hist. 343: 1. Winston, J. E. 1999. Describing Species: Practical Taxonomic Procedure for Biologists. Columbia University Press, New York. Wolfheim, J. H. 1983. Primates of the World: Distribution, Abundance and Conservation. University of Washington Press, Seattle, WA. Youlatos, D. 1999. Comparative locomotion of six sym patric primates in Ecuador. Annls. Sci. Nat. 20: 161.

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144 A I: PITHECIA r Acajutuba, Rio Negro, Amazonas, Brazil [03'S, 60'W; E. Snethlage 1916]. Acre, Estao Ecolgica do Rio Acre, Brazil [PH]. Aguarico (Ro), Sucumbios, Ecuador [-0.27, -76.38]. Altamira, Ro Manu, Madre de Dios, Peru [-11.8, -70.8 // 12'S, 71'W; C. Kalinowski, 1960, 1961]. Amacayacu National Park, Amazonas, Colombia [-3.29, -70.14]. Amazonia, Parque Nacional da; near Itaituba, Par, Brazil [-4.3. -56.7]. Amorin (Igarap), Par, Brazil [-2.0, 54.9; Olalla Brothers 1931 in Paynter Jr. and Traylor Jr. 1991]. Anavilhanas Archipelago, Estao Ecolgica, [02'S, 60'W; PH]. Andira (Lago), Amazonas, Brazil [-2.3, -56.9]. Andira (Rio), mouth; Amazonas, Brazil [-2.4, -56.8]. Aniba (Igarap); Amazonas, Brazil [02'S, 58'W; A.M. Olalla 1936]. Apuau (Rio), Amazonas, Brazil ['S, 60'W; M.C. Mello 1959]. Aripuan (old), Mato Grosso, Brazil [09'S, 60'W; F. Stolle 1914]. Aripuan (Rio) [-4.2, -60.4; F. Rhe]. Arrawarra Creek, Nickerie, Suriname [5.46, -55.71]. Arouani River, French Guiana [4.74, -53.78]. Arua, Rio Arapiuns, Par, Brazil [02'S, 55'W; A.M Olalla 1936]. Arum, Rio Purus, Amazonas, Brazil [04' S, 62'W; A.M. Olalla 1935]. Atabani (Rio) (= Itabani, Rio), Amazonas, Brazil [02'S, 58'W; A.M. Olalla 1937]. Atalaya do Norte, Rio Javari, Amazonas, Brazil [-4.24, -70.19]. Auara (Igarap), Rio Madeira, Amazonas, Brazil [4' S, 59'W; Olalla brothers. 1930]. Autazes, Rio Madeira, Amazonas, Brazil [-3.58, -59.13]. Ayapua (Lago), Amazonas, Brazil [-4.35, -62.14]. Bacabal Cuieira Apuau", Rio Negro, Amazonas, Brazil [M.C. Mello. April 1959; PH]. Baeza, Napo, Ecuador [-0.2, -77.9; W.M. Richardson, January 1913]. Balbina (Balbina Dam, Represa de Balbina, south end), Rio Uatum, Amazonas [-1.7, -59.4]. Balta, Rio Curanja, Rio Purus, Ucayali, Peru [10'S, 71'W; J.P. O'Neill 1963 and A.L. Gardner 1966, 1968, 1971 (PH)]. Barra do Rio Negro (= Manaus), Amazonas, Brazil [-3.06, -59.97]. Bartica, Demerara-Essequibo, Guyana [6.41, -58.62]. Base Atun, Ro Samiria, Reserva Nacional Pacaya-Samiria, Loreto, Peru [-5.17, -74.67]. Belen, Ro Cunucunuma, Amazonas, Venezuela [03'N, 65'W; M. Tuttle 1967]. Benjamin Constant, Rio Javari, Amazonas, Brazil [-4.25, -70.03]. Berbice; Guyana [05'N, 57'W; PH]. Bigi Poika, Para SR, Suriname [5.64, -55.49]. Blanco (Quebrada), Loreto, Peru [4'S, 73'W]. Bobonaza (Ro), Morona-Santiago, Ecuador [01'S, 77'W // mouth: -2.57, -76.63]. Boia (Lago), Rio Tef, Amazonas, Brazil [approx. 04'S, 65'W; Paynter Jr. and Traylor Jr. 1991]. Boiuu, Buiussu (Igarap) (= Boiussu, Igarap), Par, Brazil [01'S, 54'W]. Bolivia (Igarap); Amazonas, Brazil [03'S, 60'W; E. Snethlage 1916]. Bolivar Botanical Garden, Orinoco, Venezuela [7.8, -63.3]. Bomba, Ro Bomba at mouth, Loreto, Peru [07'S, 73'W; Stevens and Traylor Jr. 1983]. Bonasica (= Bonasika); Essequibo, Guyana [06'N, 58'W]. Bonnesique Creek (Bonasika River), Essequibo, Guyana [06'N, 58'W]. Borba, Rio Madeira, Amazonas, Brazil [-4.34, -59.59]. Botonamo (Rio), near Ro Cuyuni, Bolivar, Venezuela [06'N, 61'W; PH]. Branco (Rio), Acre, Brazil [-9.8, -67.8]. Branco (Rio), alto, at Rio Marac, Amap, Brazil [00'N, 52'W; PH]. Brabo, (Bravo), Par, Brazil [-2.0, 54.9; Olalla brothers 1931 in Paynter Jr. and Traylor Jr. 1991]. Bratica, Guyana [6.44, -58.61]. Bravo, Edo. do Eira (= Lara), Par, Brazil [-1.89, -55.18]. Bravo (Igarap), Rio Tapajs Par, Brazil [02'S, 55'W; PH]. Bravo Ravine, Ro Cotuhe, Amazonas, Colombia [3.03, -70.16].

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145 Brownsberg Nature Preserve, Brokopondo, Suriname [5.00, -54.87; R. A. Mittermeier]. Brownsberg Nature Preserve, Sipaliwini, Suriname [05'N, 55'W; M. Norconk]. Cacipor do Amap (Rio), Amap, Brazil [3.8, -51.2]. Cacoal Selva Park, Rondnia, Brazil [-10.9, -61.5]. Cachoeira do Tronco, Rio Erepecuru (= Rio Par do Oeste), Par, Brazil [01'S, 56'W; M. Lasso 1937]. Cahuinari (Ro), Amazonas, Colombia [-1.38, -71.65]. Caititu (Igarap), Rio Uatum, Amazonas, Brazil [-1.75, -59.75]. Canaboua, Paran do Jacar; Amazonas, Brazil [03'S, 60'W; K. Lako 1932, W. Ehrhardt 1925-26 (PH)]. Canaboua (Lago); Amazonas, Brazil [cf., Canabuoca]. Canacari (Lago) or Canacary, Amazonas, Brazil [02'S, 58'W; A.M. Olalla 1935]. Cao Agua Blanca, Curare-Los Ingleses Indigenous Reserve, Colombia [01'36"S, 69'02"W; E. Palacios] Cao Arapa, Pur National Park, Colombia [02'S, 69'W; E. Palacios] Cao Curare, Curare-Los Ingleses Indigenous Reserve, Co lombia [01'5"S, 69'22"W; E. Palacios] Cao El Boliviano, Camaritagua Indigenous Reserve / Vereda Madroo, Colombia [01'42"S, 69'58"W; E. Palacios] Cao Esperanza, Pur National Park, Colombia [01'23"S, 69'39"W; E. Palacios] Cao Mateo, Pur National Park, Colombia [02'S, 69'W; E. Palacios] Cao Zumaeta, Curare-Los Ingleses Indigenous Reserve, Colombia [01'19"S, 69'04"W; E. Palacios] Caruaro (Lago), Rio Ia at Solimes, Amazonas, Brazil [03'S, 68'W; K. Lako and Salatke (PH)]. Carvara, (Lago do), Brazil [03'S, 68'W; PH]. Cassipore (Rio) (= Rio Cacepor do Amap); Amap, Brazil [PH]. Caterpiza (Ro), Ro Santiago, Amazonas, Peru [-3.6, -77.7]. Caucaya Airport, Puerto Leguizamo, Putamayo, Colombia [-0.12, -74.76]. Caucaya (Quebrada), Limoncocha, Miguel Velasquez Farm, Putamayo, Colombia [-0.07, -74.83]. Cayenne, French Guiana [5.03, -52.33]. Cerro Azul, Ro Ucayali, Ucayali, Peru [07'S, 74'W; J.M. Schunke 1947]. Chamicuros, Ro Samiria, Loreto, Peru [05'S, 75'W; H. Bassler Collection 1927 (PH)]. Chicosa, Ro Ucayali, Ucayali, Peru [10'S, 74'W; R.W. Hendee 1927]. Cipotuba (Lago do), Rio Aripuan, Rondnia, Brazil [-4.8, -60.2]. Chupinguaia, near RO-391, Rondnia, Brazil [-12.0, -61.0]. Cobija, Ro Acre Pando, Bolivia [11'S, 68'W; C. Freese, P. Heltne, N. Castro, and G. Whiteside 1975 (PH)]. Cocha Cashu, Ro Madre de Dios, Madre de Dios, Peru [-11.88, -71.39]. Colina do Veado, bidos; Par, Brazil [01'S, 55'W; E. Snethlage 1912 (PH)]. Cononaco (Ro), Ecuador [-1.2, -75.9]. Contamana, Loreto, Peru [-7.34, -75.01]. Copataza (Ro), Pastaza, Ecuador [02'S, 77'W; C. Buckley 1877, 1878]. Coppename River, Lolopasi west bank, across from Foen goe Island, Sipaliwini, Suriname [4.86, -55.96]. Crrego do Campo, Pimento Bueno, Rondnia, Brazil [q.v.; Comisso Rondon, 1914 (PH)]. Cotuhe (Ro), Tarapaca, Amazonas, Colombia [-2.77, -69.72]. Cravari, Mato Grosso, Brazil [PH]. Cruzeiro do Sul, Amazonas, Brazil [-7.63, -72.68]. Cuieiras (Rio), mouth of,Rio Negro, Amazonas, Brazil [-2.83, -60.50]. Cuipeua (Lago), near Boiussu, Par, Brazil [PH]. Cuiteua (Lago) (= Lago Cuipeua), Par, Brazil [-1.84, -55.5]. Cumeria (= Cumaria), Ro Ucayali, Ucayali, Peru [09'S, 74'W; R.W. Hendee, August 1927]. Curaca (Quebrada), Loreto, Peru [04'S, 73'W; F. Cornejo]. Curaray (Ro), mouth; Loreto, Peru [-2.0, -74.1]. Curaray and Nahio (between ros), Peru [-1.3, -75.5; R. Aquino]. Curaray (Ro), mouth, at Ro Napo, Ecuador [02'S, 74'W; Olalla brothers, October, November, Decem ber 1925]. Curumo (Ro), Bolivar, Venezuela [07'N, 61'W; PH]. Cuyabeno National Park (incl. Laguna Garza Cocha) [-0.10, -75.86]. Dadanawa, Rupununi, Guyana [02'N, 59'W; PH]. Demerara Coast Region, Guyana [7.5, -58.6]. Demerara River, Guyana [5.88, -58.4].

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146 Destacamento, Ro Maraon, Loreto, Peru [03'S, 72' W; M. Ximenez de la Espada, 1865]. Destacamento do Ribeiro, Rondnia, Brazil [-9.6, -65.3; Natterer 1883] Ega (= Tef), Amazonas, Brazil [cf. Tef] Eiru (Rio), mouth at Rio Juru, Amazonas, Brazil [06'S, 69'W]. Eirunep (= Joo Pessoa); Amazonas, Brazil [06'S, 69'W]. El Dorado, Bolivar, Venezuela [06 44'N, 61 38'W; PH]. El Hacha (Quebrada) Caquet River Expedition 1960 [-0.15, -74.57]. El Inerno Farm, Puerto Santander, Ro Caquet, Caquet, Colombia [0.70, -75.20]. El Manaco, 59 Km SE of El Dorado at Km 74, Bolivar, Venezuela [7.2, -61.6]. Erepecur (Rio) (now upper Rio Par do Oeste) [01'N, 56'W]. Espelho (Serra), Par, Brazil [cf. Faro, PH]. Esperanza, Quebrada, Ro Yavari-Mirim, Loreto, Peru [04'S, 71'W; C. Kalinowski, 1957]. Essequibo (River), West Demerara, Guyana [6.70, -58.58]. Estacin Biolgica Pithecia Loreto, Ro Samiria, Peru [05'S, 74'W; PH]. Estacin Biolgica Panguana, Rio Llullapichis, Huanaco, Peru [09'S, 74'W]. Estribaciones, Ecuador [00'S, 77'W; L. Albuja]. Estiro do Equador, Ro Javari, Amazonas, Brazil [-4.38, -71.55]. Faro, Rio Nhamunda, Par, Brazil [02'S, 56'W]. Flexal (Lago) (also see Bravo), Par, Brazil [-1.89, -55.18]. Fonte Boa, Rio Solimes, Amazonas, Brazil [-2.44, -66.07]. Fordlndia, Rio Tapajs, Par, Brazil [03'S, 55'W; PH]. Forte do Rio Branco, Roraima, Brazil [03'N, 60'W]. Galvez (Ro), Loreto, Peru [05' S, 73'W]. Gi-Paran (Rio), (= likely Rio Ji-Paran), Rondnia, Brazil [08'S, 62'W; PH]. Gordo (Igarap), Rio Juru, Amazonas, Brazil [06'S, 69'W; A.M. Olalla, 1936]. Grande (Igarap), left bank Rio Juru, Amazonas, Brazil [06'S, 69'W; A.M. Olalla, 1937]. Grande (Lago), Rio Juru, Amazonas, Brazil [06'S, 69'W; A.M. Olalla 1936]. Grande (Ro), Bolivar, Venezuela [08'N, 61'W; PH]. Guajar-Mirim, Rondnia, Brazil [-10.80, -65.34]. Guapor (Rio); Rondnia, Brazil (= Ro Mamor, Beni, Bolivia) [11'S, 65'W]. Guri, Operacin Rescate, Ro Caroni, Bolivar, Venezuela [07'N, 63'W; PH]. Guri (Lago), Bolivar, Venezuela [8.2, -62.9]. Gurisoco, El Palmar; Bolivar, Venezuela [08'N, 61'W; PH]. Hanover, Weg (nearby), Para SR, Suriname [5.57, -55.13]. Humait, Rio Madeira, Amazonas, Brazil [-6.9, -63.0]. Humboldt, Von (= Von Humboldt), Huanuco, Peru [08'S, 75'W; PH]. Iaunari (= Yavanari or Javanari), Amazonas, Brazil [-0.3, -64.8]. Itaboca, Rio Purus, Amazonas, Brazil [04'S, 62'W; A.M. Olalla 1935]. Inuya (Ro), mouth at Rio Urubamba, Ucayali, Peru [-10.68, -73.46]. Ipousin (= Ipoucin Crique), French Guiana [04'N, 52'W]. Iqu-Aripuan, Estao Ecolgica, Mato Grosso, Brazil [PH]. Iqu, Estao Ecolgica, Mato Grosso, Brazil [PH]. Iqu-Juruena, Estao Ecolgica, 11''S, 58' 59' W; E.Z.F. Setz and K. Milton 1983 (PH)]. Iquitos, Ro Amazonas, Loreto, Peru (03'S, 73'W). Iranduba, Rio Amazonas, Amazonas, Brazil [-3.0, -60.2]. Itabani (Rio), or Atabani, Amazonas, Brazil [02'S, 58'W; A.M. Olalla 1937]. Itaboca, Rio Purus, Amazonas, Brazil [-4.83, -62.76]. Itacoatiara, Rio Amazonas, Amazonas, Brazil [-3.04, -58.44]. Itaituba; Par, Brazil [04'S, 55'W; PH]. Jaburu, Rio Purus, Amazonas, Brazil [05'S, 64'W; PH]. Jacaracanga; Par, Brazil [-6.23, -57.77]. Jamund (cf. Nhamund), Par, Brazil. Jari (Rio) at Lago Jari, Par, Brazil [-4.93, -62.37]. Jaru, Reserva Biolgica Federal; Rondnia, Brazil [10' S, 62'W; PH]. Javanari, (cf. Yavanari), Amazonas, Brazil. Joo Pessoa (= Eirunep), Amazonas, Brazil [06'S, 69'W; A.M. Olalla, 1936]. Juma Lodge, Lago Juma/Rio Juma, Amazonas, Brazil [03'S, 59'W]. Juru (Rio); mouth, at Rio Amazonas, Amazonas, Brazil [-2.64, -65.75]. Kabalebo River, Sipaliwini, Suriname [4.40, -56.98].

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147 Kaieteur Falls, Essequibo, Guyana [5.27, -59.49]. Kaiserberg Airstrip; Nickerie, Suriname [03'N, 56'W; PH]. Kapoeri Creek, Nickerie, Suriname [5.41, -57.13]. Kayser Gebergte, current airstrip nearby, Sipaliwini, Suri name [3.3, -56.4]. Kayser Mountains, Sipaliwini, Suriname [3.3, -56.7; M. van Roosmalen study]. Kalacoon, Mazaruni-Potaro, Guyana [06'N, 58'W; Stephens and Traylor Jr. 1985]. Kartabo, Mazaruni-Potaro, Guyana [06'N, 58'W; Stephens and Traylor Jr. 1985]. Km 16, BR 230, Rondnia, Brazil [-7.2, -62.6]. Km 19, Itaituba-Jacaracanga, Par, Brazil [06'S, 57'W; PH]. Km 21, BR 174 Tarum, Manaus, Amazonas, Brazil [-2.97, -60.0]. Km 104-200, Manaus-Itacoatiara; Amazonas, Brazil [02'S, 58'W; M. Mello 1967]. Km 105, BR 174 N, Manaus, Amazonas, Brazil [-1.94, -60.08]. Km 212, Itaituba-Jacaracanga, Para, Brazil [04 53' S, 56 47' W; PH]. Km 165, Manaus-Itacoatiara; Amazonas, Brazil [03'S, 59'W; M. Mello 1965, A. Faustino 1965]. Km 170, Manaus-Itacoatiara; Amazonas, Brazil [03'S, 58'W; M. Mello 1967]. Km 190, Manaus-Itacoatiara; Amazonas, Brazil [02'S, 58'W; M. Moreira 1966, M. Mello 1967]. Kourou Road, Saut Laurent, French Guiana [5.38, -53.07]. La Coca, Ro Napo, Napo, Ecuador [00'S, 76'W; M. Jimnez de la Espada, 1865]. Lagarto, Ro Ucayali, approx. Olalla camp, Ucayali, Peru [-10.64, -73.81]. Lagarto Cocha (mouth), border Ecuador/Peru [00'S, 75'W; Olalla brothers 1926 1928]. Lago Piratuba, (Reserva Ecolgica), Amap, Brazil [01'N, 50'W; A. B. Rylands and R. A. Mittermeier 1982 (PH)]. La Chorrera, Igara-Paran (Rio), 20 km downstream of La Raicilla Ravine, Amazonas, Colombia [-1.47, -72.62]. La Macarena, Meta, Colombia [2.21, -73.94]. La Muralla (= Morelia), Caquet, Colombia [1.50, -75.73]. La Poule, Saramacca, Suriname [05'N, 55 25'W; PH]. Las Patas, Bolivar, Venezuela [07'N, 62'W; PH]. La Solita Creek, Solita, Colombia [00'N, 75'W]. La Trinidad, El Palmar, Bolivar, Venezuela [07'N, 61'W; PH]. Lely Gebergte, Sipaliwini, Suriname [4.47, -54.65]. Limoal, near Boim, Par, Brazil [-2.82, -55.17; Paynter Jr. and Traylor Jr. 1991]. Limoatuba, Par, Brazil [-2.0, 54.9; Olalla brothers 1931 in Paynter Jr. and Traylor Jr. 1991]. Limontuba (= Limoatuba), Par, Brazil [PH]. Llullapichis (= Yuyapichis) (Ro) mouth, Huanuco, Peru [-9.62, -74.96]. Loksie Hattie, Suramaca River, Brokopondo, Suriname [5.33, -55.35]. Los Amigos Concesin para Conservacin, Ro Los Amigos, Madre de Dios, Peru [12'S, 70'W]. Macap, Amap, Brazil [0.03, -51.07]. Macas, Morona-Santiago, Ecuador [-2.1, -78.1]. Maisisa, Ucayali, Peru [08'S, 74'W; Stevens and Traylor Jr. 1983]. Mamirau Reserve, Amazonas, Brazil [-2.40, -65.36]. Mamore (Ro), mouth, Beni, Bolivia, also Rio Guapor, Rondnia, Brazil [-10.43, -65.39]. Manacapur, Rio Solimes, Amazonas, Brazil [-3.1, -60.6]. Manaquiri, south of Rio Amazonas, Amazonas, Brazil [-3.37, -60.42]. Manarica Creek, Essequibo River, Guyana [PH]. Manaus, Amazonas, Brazil [-3.06, -59.97]. Manu, Parque Nacional, Madre de Dios, Peru [-11.8, -70.8]. Mapixi (Lago), (= Lago Mapiocu), Rio Purus, Amazonas, Brazil [05'S, 63'W; K. Lako 1931]. Marac (Rio), Roraima, Brazil [-0.1, -51.6]. Mararu, Par, Brazil [-10.43, -65.39]. Marua (= cf. Mararu); Par, Brazil. Matapi Creek and Kabalebo River near Corantijn River, Suriname [5.08, -57.60]. Matapi, Suriname [5.11, -57.30]. Mato Grosso, Mato Grosso do Sul (Natterer 1826), Brazil [-14.3, -59.9]. Mazaruni River, Essequibo, Guyana [06'N, 58'W; PH]. Mazan, Ro Amazonas, Loreto, Peru [-3.47, -73.06]. Mecaya (Ro), below mouth, right bank, Ro Caquet, Pu tumayo, Colombia [00'N, 75'W; P. Hershkovitz, March 1952]. Miguel (Lago), (approx. now Lago Bonm), Rio Madeira, Amazonas, Brazil [-3.79, -59.11]. Mocoa, Putamayo, Colombia [1.26, -76.65]. Moengo, SW, Suriname [5.66, -54.44].

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148 Moengotapoe, Marowijne, Suriname [5.65, -54.29]. Montalvo, Ro Bobonaza, Pastaza, Ecuador [02'S, 76'W]. Montanita, Florencia, Caquet, Colombia [01'N, 75'W]. Montealegre (= Monte Alegre), Huanuco, Peru [09'S, 74'W; Tessman, November 1922]. Morelia, Ro Fragua, Caquet, Colombia [01'N, 75'W; L.E. Miller, July 1912]. Moroni River, Suriname [5.52, -54.05]. Mucden (= Mukden), Pando, Bolivia [-11.0, -69.1], Mukden, Pando Bolivia [11'S, 69'W; L. Porter, eld site] Napo Wildlife Center, Ro Napo, Ecuador [-0.2, -76.8]. Napo (Ro) at Coca, Orellana, Ecuador [-0.46, -76.98]. Napo (Ro) north near mouth of Ro Curaray, Loreto, Peru [-22.0, -74.0; Olalla brothers collection 1925]. Napo (Ro), north of Iquitos, Loreto, Peru [-1.57, -74.21; Olalla brothers collection 1925]. Nareuda (Ro), Mukden, Bolivia [11'S, 6'W]. Nashio (Ro), Peru [near -.98, -75.31; R. Aquino]. Nassau Gebergte, near Marowijne River, Sipaliwini, Suri name [4.85, -54.60]. Nauta, Loreto, Peru [-4.58, -73.77]. Navio, Serra do (=Vila do Serra), Amap, Brazil [1.1, -52.0]. Negro (Rio) mouth at Rio Amazonas, Amazonas, Brazil [-3.13, -59.90]. Nevati (Mission, right bank), Ro Pichis, Oxapampa, Pasco, Peru [10'S, 74'W; D.R. Seidel 1964, M.D. Tuttle 1963]. Nhamund (Rio), Par, Brazil [02'S, 56'W]. Nova Braslia, Rondnia, Brazil [-10.4, -61.9]. Nuevo San Juan, Ro Galvez, Loreto, Peru [05'S, 73'W]. Nuria (Altiplanicia); Bolivar, Venezuela [07'N, 61'W; PH]. bidos, Par, Brazil [-1.84, -55.51]. Oiapoque, French Guiana [3.97, -51.83]. Oko Mountains, Cuyuni-Mazaruni, Guyana [06'N, 58'W; E.R. Blake 1937 (PH)]. Oriximin; Par, Brazil [-1.4, -55.9]. Orosa, Ro Amazonas, Loreto, Peru [03'S, 72'W; Olalla brothers 1926]. Orosa (Ro), Loreto, Peru [-3.69, -72.44; R. Aquino]. Pacas Novos (Rio), Rondnia, Brazil [-10.85, -65.00; Natterer 1829]. Pacas Novos, Parque Nacional, Rondnia, Brazil [10'S, 63'W; PH]. Pacaya-Samiria, Reserva Nacional, Loreto, Peru [-5.20, -74.59]. Pachitea (Ro) mouth, Huanuco, Peru [-8.74, -74.50]. Palmares, Amazonas, Brazil [PH]. Palomeu River Camp, Tapanahoni River, Marowijne, Suri name [3.3, -55.4]. Papel Azul (Igarap); Amazonas, Brazil [02'S, 60'W; PH]. Paramaribo, Suriname [5.87, -55.16]. Pastaza (Ro) mouth at Rio Maraon, Pastaza, Peru [04'S, 76'W]. Paranova (Lagoinho), Rio Guapor, Rondnia, Brazil [11'S, 65' W; J. Natterer, 1829]. Paratuco (Rio) (= Piratucu), Ama zonas, Brazil [01'S, 56' W; Olalla brothers 1930]. Perica River, Moengo, Suriname [5.91, -54.69]. Parinari (= Parinari Cao?), Loreto, Peru [-4.1, -74.5; Bluntsali-Peyer Expedition 1913]. Parintins, Amazonas, Brazil [-2.2, -56.7]. Parque Nacional da Amaznia, Rio Tapajs, Par, Brazil [03'S,57'W; PH]. Par do Oeste (Rio), Par, Brazil [-1.1, -56.0]. Passarinho (Igarap), Amazonas, Brazil [PH]. Patana (Cachoeira), Par, Brazil [3.13, -62.35]. Pebas, Ro Amazonas, Loreto, Peru [03'S, 71'W; R.W. Hendee 1928]. Penal Settlement, Burntbush, Mazaruni-Potaro, Guyana [06'N, 58'W; PH]. Piratuba, Lago; Amap, Brazil [01'N, 50'W; A. B. Rylands and R. Mittermeier 1982 (PH)]. Pisqui (Ro), Ucayali, Peru [07'S, 75'W; H. Bassler Collection, 1923].23 Ponta da Castanha, SW bank Rio Tef, Amazonas, Brazil [-3.81, -64.97 approx; A. Johns 1985]. Porongaba, Rio Juru, Acre, Brazil [-8.66, -72.82]. Porteira, Par, Brazil [-0.8, -57.0]. Porto Mau, Manaus, Amazonas, Brazil [03'S, 60'W; M. Oreira 1967, F. Almeida 1965]. Porto Saids, Rio Juru, Acre, Brazil [-8.17, -72.76]. Porto Trios, Par do Oeste, Par, Brazil [02'N, 55'W; PH]. Potaro Landing, Essequibo, Guyana [05'N, 59'W, PH]. Pucallpa, Ucayali, Peru [-8.35, -74.55].

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149 Puerto Indiana, Ro Amazonas, Loreto, Peru [-3.4, -73.04 //03'S, 73'W; H. Bluntschli, 1912; Olalla brothers, 1926]. Puerto Japn, Ro Peneya, Caquet, Colombia [00'N, 74'W]. Puerto Leguizamo and La Tagua [between rivers: -0.05, -74.73; K. Watanabe Expedition 1973]. Puerto Melendez, N bank Rio Maraon, Loreto, Peru [04"S, 77'W; H. Bassler Collection, February 1929]. Puerto Pachitea, Ro Pachitea, Huanaco, Peru [-9.93, -75.03]. Puerto Pungo, Ro Punga, Loreto, Peru [-5.7, -74.1]. Puerto Pungo, Ro Tapiche, Loreto, Peru [06'S, 74'W; PH]. Puerto Tokio, Ro Peneya, Caquet, Colombia [00'N, 74'W]. Puerto Victoria, Ro Pachitea, Huanaco, Peru [-9.88, -74.94]. Quebradn El Ayo, Pur National Park, Colombia [01'S, 69'W; E. Palacios] Quixito (Ro), Loreto, Peru [04'S, 70'W]. Reserva de Desenvolvimento Sustentavel Piaga-Purus, Rio Purus, Amazonas, Brazil [-4.5, -62.3]. Reserva Nacional Allpahuayo-Mishana, Loreto, Peru [-3.91, -73.12]. Reserva Nacional Pacaya-Samiri, Loreto, Peru [05'S, 74'W] Retiro do Veado Branco, Serra do Norte, Rondnia, Brazil [11'S, 59'W; Comisso Rondon]. Rio Acre, Estao Ecolgica, Acre, Brazil [10'S, 70'W; PH]. Rockstone, Essequibo River, Guyana [5.98, -58.53]. Rosarinho, Amazonas, Brazil [03'S, 59'W; Olalla brothers 1930]. Rupununi River, mouth, Rupununi, Guyana [4.0, -59.0]. Samiria (Ro), mouth; Loreto, Peru [04'S, 74'W]. Sampaio (Lago), Rio Madeira, Amazonas, Brazil [-3.84, -59.17]. San Fernando, Ro Yavari-Mirim, Loreto, Peru [04'S, 70'W; C. Kalinowski, 1957]. San Francisco, left bank Ro Napo, Napo, Ecuador [00'S, 76'W; P. Hershkovitz, March 1936]. San Jose del Encanto, Ro Caraparana, Putumayo, Colom bia [01'S, 73'W; H. Hill, 1975, 1976]. San Martn de Turumbari, Ro Cuyuni, Bolivar, Venezuela [06'N, 61'W; PH]. San Lorenzo, near border with Bolivia, Madre de Dios, Peru [-11.3, -69.3; R. Aquino Collection]. San Salvador, Ro Tapiche, Amazonas, Peru [-5.0, -73.9]. Santa Cecilia, Ro Maniti, Loreto, Peru [03'S, 72'W; C. Kalinowski, June 1957]. Santa Cruz, Rio Eiru, Amazonas, Brazil [07'S, 70'W; PH]. Santa Elena, Ro Samiria, Loreto, Peru (05'S, 74'W; PH). Santa Lucia (not Santa Luisa), Ro Nanay, Loreto, Peru [03'S, 74'W; C. Kalinowski 1956 (PH)]. Santa Luisa (likely location), Ro Nanay (lower near Rio Maraon), Loreto, Peru [-4.35, -74.12; Stephens and Traylor Jr. 1983]. Santarm, Rio Tapajs, Par, Brazil [-2.55, -54.72]. Santa Rita do Weil (= Santa Rita), Rio Solimes, Amazo nas, Brazil [-3.48, -69.31; W. Ehrhardt, 1926]. Santiago (Ro) mouth, Esmeraldas, Ecuador [-4.30, -77.63]. Santo Antnio, Rio Eiru, Amazonas, Brazil [06'S, 69'W; PH]. Sapote (Quebrada), Ro Ucayali, Loreto, Peru [-5.18, -74.15; R. Snyder]. Sarayacu, Ro Ucayali, Loreto, Peru [-6.72, -75.10]. Sarayaku, Morona-Santiago, Ecuador [-1.6, -77.5]. Saut Laurent du Maroni, French Guiana [5.60, -53.99]. Saut Sabbat, French Guiana [5.50, -53.66]. Saut Tigre, French Guiana [5.38, -52.93]. So Felipe at or below, Rio Juru, Amazonas, Brazil [unknown location, ca. 07S (PH); E. Garbe, 1901-1902]. So Joaquim, Rio Branco, Roraima, Brazil [03'N, 60'W; PH]. So Manoel (= So Manuel), Rio Teles Pires, Par, Brazil [07'S, 58'W; F. Hoehne 1912, 1914]. So Paulo (Fazenda), Amazonas, Brazil [q.v.]. So Sebastio do Uatum, Rio Uatum, Amazonas, Brazil [-2.56, -58.15]. Saramacca, Suriname [6.5, -55.5]. Sarayacu, Pastaza, Ecuador [-1.5, -77.5]. Sarayacu, Ro Bobonaza, Pastaza, Ecuador [01'S, 77'W; PH]. Sarayacu, Ucayali, Peru [-6.4, -75.0]. Sarayacu, Olalla Camp (March-May 1927), Ucayali, Peru [-6.63, -75.04; Stevens and Traylor Jr. 1983]. Sarayacu, Olalla Camp (May-June 1927), Ucayali, Peru [-6.59, -75.14; Stevens and Traylor Jr. 1983]. Sarayacu, Olalla Camp (August 1927), Ucayali, Peru [-6.89, -75.10; Stevens and Traylor Jr. 1983]. Seringal Oriente, Rio Juru, Acre, Brazil [08'S, 72'W; PH].

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150 Serpa (Lago), Itacoatiera, Amazonas, Brazil [03'S, 58'W; A. M. Olalla 1936-1937]. Serra de Espelho, Faro, Par, Brazil [02'S, 56'W; A. M. Olalla 1930, 1937]. Shiripuno Research Lodge, Ro Shiripuno, Ecuador [01'S, 76W; L. K. Marsh]. Sushundi, Succumbios, Ecuador [-0.04, -76.64]. Siguin (or Sihuin), Ro Pastaza, Loreto, Peru [03'S, 76'W; C. Olalla, Mission Flornoy, 1936]. Silves, Amazonas, Brazil [02'S, 58'W; A. M. Olalla 1937]. Sipaliwini Airstrip (1970s), Sipaliwini, Suriname [3.9, -56.2]. Supamo (Ro), Bolivar, Venezuela [07'N, 62'W; PH]. Supinaam River, Demerara, Guyana [06'N, 58'W; Stephens and Traylor Jr. 1985]. Tabatinga, Amazonas, Brazil [-3.7, -69.6]. Tabalosos, Huanuco, Peru [-6.36, -76.63]. Tabocal (Tabucal), Amazonas, Brazil [-3.03, -58.18 // Tabucal: 00 47' S, 67 15' W; A. M. Olalla 1929]. Tacshacurary (Rio) at Ro Machete (mouth) [-2.66, -73.62]. Tahuamanu (Ro), Pando, Bolivia [11'S, 68'W; K. Izawa 1978]. Tahuayo Lodge, Ro Tahuayo, Loreto, Peru [-4.22, -73.22]. Tamashiyacu, Ro Amazonas, Amazonas, Peru [-4.01, -73.15]. Tamboryacu (Ro), Loreto, Peru [-1.5, -74.2; R. Aquino]. Tapanahoni River (cf., Paloemeu River) [3.3, -55.4]. Tapiche (Ro), Amazonas, Peru [-5.0, -73.9]. Tapirapoan (= Tapirapua); Mato Grosso, Brazil [PH]. Tapirapua, Rio Cipotuba, Mato Grosso, Brazil [14'S, 57'W; F. Hoehne 1914]. Taumaturgo (Vila), Acre, Brazil [PH]. Tef (Lago), Amazonas, Brazil [03'S, 64'W; H. Bates 1850-54 (PH)]. Tef (Rio); Amazonas, Brazil [-3.81, -64.97]. Teresinha, Rio Amapar, Amap, Brazil [00'N, 52'W; PH]. Terra Santa, Lago do Algodal, Par, Brazil [02'S, 56'W; PH]. Tigrillo (Ro), Ro Tigre, Peru [04'S, 74'W; PH]. Tiputini Biodiversity Station, Ro Tiputini, Yasun Nation al Park, Ecuador [-0.5, -76.3]. Trios, Rio Par do Oeste, Par, Brazil [02'N, 55'W; PH]. Tonantins (Rio), Amazonas, Brazil [-2.79, -67.87; J. B. von Spix 1820]. Tracajatuba (Rio), Amap, Brazil [1.1, -51.1]. Tres Troncos, La Tagua, Ro Caquet, Colombia [00'N, 74'W]. Trombetas, Rio, Reserva Biolgica, Par, Brazil [01'S, 57'W; A. B. Rylands and R. A. Mittermeier 1982]. Tumatumari, Potaro River, Essequibo, Guyana [05'N, 59'W; PH]. Tushemo (= Tushma or Tusma), Ucayali, Peru [08' S, 74'W; L. Rutter, 1926]. Tutapisco (Ro), Peru [-3.02, -73.26]. Uarin, Par, Brazil [-2.91, -65.10]. Uatum (Rio), Amazonas, Brazil [-2.56, -58.15]. Uaua (Lago), (Coar), Amazonas, Brazil [-4.19, -62.37]. Urubamba (Ro), mouth at Ro Ucayali, Ucayali, Peru [10'S, 73'W; Olalla brothers, 1927]. Urubamba (Ro) lower, T. Gregory Field Site, Ucayali, Peru [-11.74, -72.91]. Urub (Rio), mouth, Amazonas, Brazil [02'S, 58'W; PH]. Uruc (Rio), Amazonas, Brazil [-4.46, -64.35]. Utoquinea; Ucayali, Peru [-9.67, -74.08 // 08'S, 74'W; H. Bassler collection 1923]. Vereda La Leona, Valparaiso, Colombia [01'N, 75'W]. Vila Bella Imperatriz, Lago Andira, Amazonas, Brazil [02'S, 56'W; Olalla brothers 1930]. Villa Garzn, Puerto Limn, El Picudo Indigenous Reser vation, Putumayo, Colombia [1.13, -76.60]. Vila Braga, Par, Brazil [04'S, 56'W. [F. Lima, 1917]. Vila Nova (Rio), Amap, Brazil [00'N, 51'W; PH]. Voltzberg, east bank, Coppename River, Sipaliwini, Suriname [4.81, -56.04]. West River, Wilhelmina Mountains, Nickerie, Suriname [03'N, 56'W; PH]. Wia-Wia Nature Preserve, Moengo, Suriname [5.94, -54.44]. Wilhelmina Mountains, approx. West River, Sipaliwini, Suriname [4.0, -56.6]. Xapur, Acre, Brazil [-10.4, -68.5]. Yanayacu, Ro Maraon, Loreto, Peru [-4.79, -74.95]. Yarina Eco Lodge, Ro Napo, Napo, Ecuador [-0.3, -76.8]. Yavanari (= Iaunari); Amazonas, Brazil [-0.3, -64.8]. Yavari (Ro), near mouth, Loreto, Peru [04'S, 70'W; PH]. Yavari Mirim (Ro), Loreto, Peru [-4.51, -72.70]. Yavari Mirim (Ro) mouth, Loreto, Peru [04'S, 71'W].

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151 Yotacuay; Delta Amacuro, Venezuela [08'N, 61'W; PH]. Yuturi Eco Lodge, Ro Napo, Ecuador [-0.5, -76.0]. Zanderij, Para SR, Suriname [5.60, -55.21]. Zinho (Igarap) near Igarap Aniba, Amazonas, Brazil [-2.91, -58.55]. Zuid River, Sipaliwini, Suriname [3.4, -56.4]. Unspecied coordinates per Gmap. ^ PH indicates locations reported by P. Hershkovitz (1987).

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152 A II. T Jr S Introductione specimens presented in this appendix were mostly collected by H. Schneider in the inundation area of the Samuel hydroelectric dam (Jirau dam) of the Rio Jamari, Rondnia, Brazil. ese animals appear distinct, particu larly the females which do not resemble any other females in the genus, at least by the specimens available. While it was tempting to declare them as a distinct species, more data needs to be collected on living sakis in that region. e information below follows a taxonomic format, but is without types or names.Synonymy 1987 P. irrorata Hershkovitz, P. (1987). Am. J. Primatol. 12: 425, in part. Key Specimen I MPEG No. 21961, adult male, skin and skull, collected by H. Schneider and E. Martins, 29 April 1988. Key Specimen II. MPEG No. 21785, adult male, skin and skull, collected by H. Schneider, 2 February 1988. Additional Important Specimens. Males: MPEG Nos. 21775 subadult, skin (H. Schneider, 7 December 1987); 21791 subadult, skin (H. Schneider, 25 March 1988); 21792 subadult, skin and skull (H. Schneider, 3 March 1988); 21937 subadult, skin and skull (H. Schneider, 29 August 1987); and 21959 adult skin (H. Schneider and E. Martins, 1987). Females: MPEG collection Nos.: 21787 adult, skin and skull (H. Schneider, 18 January 1988); 21784 adult, skin and skull (H. Schneider, 2 February 1988); 21778 adult, lactating, skin and skull (A.P. de Souza Jnior and F. Braga, 19 November 1987); 21782 adult, skin and skull (A.P. de Souza Jnior and F. Braga, 23 November 1987); 21934 adult, skin and skull (A.P. de Souza Jnior and F. Braga, 27 November 1987); 21960 adult, skin and skull (H. Schneider and E. Martins, 1987). MNRJ Nos.: 28479 adult, skin with 28480 infant male, skin and skull (A.P. de Souza Jnior and E. Dente, 24 January 1988). Key Specimens Locality. All key specimens from Samuel hydroelectric dam, both sides of Rio Jamari, upper Rio Madeira right side, Rondnia, Brazil. Total Specimens Examined 32 skins and skulls. I have not seen any conrmed living photos. Description. Males. All males, young and old, have the front half of their back legs black (to varying degrees in young animals), and very black wrist cufs on the forearms in all ages. Based on the type series, adult males can be irrorata-like or rylandsi-like. e rylandsi-like male colorations (MPEG No. 21961) are very grizzled white with very black back legs, but with orange rufs more simi lar to either P. irrorata or P. mittermeieri. Hands and feet are of-white. Faces are primarily black, but can be dark red throughout with little facial hair. ere maybe a few stray white hairs between the eyes, but any facial hair is con centrated to the very white stif lip and lower nose hairs, without discernable muzzle linessimilar to P. rylandsi. Older males may be like either rylandsi or mittermeieri in that they can get very white. A very white male with an orange ruf MPEG No. 21959, skin only, is an example, but the face is not preserved well. e irrorata-like adult males are less grizzled, have rufs with dark roots and light tips, have distinct white brow bands, and have black muz zles with unpigmented or reddish facial skin. ey have light, but distinct muzzle lines in addition to the stif white lip hair. In MPEG No. 21785, there are some light white hairs above the eyes as well. Subadult and juvenile males are less and more uniformly grizzled throughout their dorsal pelage. eir rufs are varied depending on age from very light bufy to darker with light tan roots to more orange. Young faces are mottled pink and black, but do not have distinct eyespots as in P. rylandsi, the pattern is much more difuse. It is possible that as they age they become more irrorata-like but it is unclear for how long as older skulls belonging to both the black faces and the irrorata faces are among the collection. Females. Adult females can look similar to the irroratalike males in that they have black muzzles and unpigmented or reddish facial skin with white brow bangs. ey have lighter eyespots of similar color to most of the face, save where there are patchy black pigments or the black ish muzzle. Even in young animals they have shaggy white malar lines and white lips that are not as distinct as the males. ey have some light black facial hair surrounding the face in subadult females, similar to P. irrorata females. ey also have white eyebrows similar to P. rylandsi subadult females, but in most cases it appears as more of a star in the forehead between the eyes. In older females, this brow can be more concentrated to between the eyes with light white and black hair dispersed over the rest of the face or a light trace of white that covers the centerline of the forehead from the hairline to the eyes. e rest of the pelage is uniformly moderately grizzled with forearms densely covered in white to bufy hairs. Rufs are less de ned than males, are dark brown, and are nely haired. Faces are variable in that when young they are very pink and unpigmented, but might have stif white hairs forming as the eyebrows. Subadult to adult females have an enlarged vulva and obvious clitoris, and can easily be mistaken for

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153 males especially in dried specimens. Infant females look just like males in preserved specimens. Measurements: Tables 27 and 28. Distribution. Map 18. Mostly collected in the region of the Samuel hydroelectric dam on the Rio Jamari, both sides of the river. It is possible that these sakis occur in the entire region of the Mamor-Madeira-Jamari. Discussion. Specimens from MPEG and MNRJ were collected by H. Schneider, and A.P. de Souza Jnior and F. Braga between 1987 and 1989. A juvenile male collected by S.F. Ferrari on 5 September 1990 along the BR 230 (Humait-Itaituba) at km 16 appears to belong to this group. If they do prove to be distinct, it would extend their distribution much further north. However, it is entirely possible that this (and possibly all) Jamari sakis belong to P. mittermeieri which dominates the region. e Jamari sakis are an interesting group primarily because adult males can resemble both a variation of P. irrorata and P. mittermeieri males (although the Jamari animals in the black phase seem like a cross between P. mittermeieri and P. rylandsi). It is unclear if these animals are hybrids, if they are polymorphic, or if they are a described species that is very long-lived and continue to transition colors, such as P. mittermeieri or P. rylandsi. Likewise, it will be important to study in detail P. irrorata populations, particularly in the state of Acre to determine boundaries with this Jamari group, and with P. mittermeieri and P. rylandsi populations north of Rio Guapor. Looking at females alone as compared to the species potentially in the areairrorata, mittermeieri, rylandsithe Jamari saki females are distinct as adults. And in general, it appears Jamari sakis are unique when the skulls are compared, but more study is needed. ere are specimens in the Comisso Rondon collection at MNRJ that are similar to some of the sakis in the Jamari collection (for example, MNRJ No. 3335 and MPEG No. 21937). Comparing both collections, however, does not yield enough similarities, although the possibility is not ruled out that they are the same as those collected in the upper Aripuan region during the Comisso Rondon ex peditions; however, it is equally likely that the Comisso Rondon animals are legacy P. irrorata (cf. P. irrorata) from a long ago larger distribution of these animals. It is also entirely possible that these animals are P. mittermeieri, but with a pelage change over the decades since the Comisso Rondon. Because of the extreme variation in these specimens, it is dicult to say if they have ever been photographed living. It is possible that if females had darker faces and white eye brows only, they might look similar to P. rylandsi subadult females, and likewise that subadult males may be mistaken for either P. irrorata, P. rylandsi or P. mittermeieri especially in captivity. It is critical to get some photos or to observe living sakis in the Rio Jamari watershed, particularly around Samuel hydroelectric dam, as they are likely in highly frag mented habitat, and are threatened no matter the ultimate species determination. Table 27. Post-cranial measurements (mm) and weights (g) of specimens. Specimen Total Length Tail Hind Foot Ear Weight MNRJ 28479 AF 860 460 110 32 1550 28480 INF 460 260 75 335 MPEG 21776 SAF 750 450 112 17 1100 21777 SAF 760 450 119 26 1100 21783 JM 660 360 107 34 930 21784 AF 833 435 116 35 1780 21785 AM 860 440 116 36 2750 21786 AF 785 450 115 30 1700 21787 AF 800 410 115 31 1800 21788 SAF 760 385 110 29 2010* 21789 INF 550 310 93 13 560 21790 JF 730 380 105 28 1280 21934 AF 840 440 116 30 2000 21935 JM 665 380 95 31 800 21936 INF 450 230 74 25 300 22001 JM 734 402 114 33 1580AM = adult male, SAM = subadult male, JM = Juvenile male, AF = adult female, SAF = subadult female, JF = juvenile female, INF = infant. *Pregnant?

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154 Table 28. Skull measurements (mm) from the Hocrcio Schneider and Arlindo Pinto de Souza Jnior and Fernando Braga collections at MPEG. Specimen MPEG Nos. Length of brain case Width of brain case Zygomatic arch width Orbits-outer Nose bridge Lorbit inner width Lorbit inner height Muzzle width Lmandible length Lmandible height Males 21785 87.0 44.1 59.6 47.2 8.7 17.1 18.4 25.9 59.7 40.5 21783 72.9 44.1 43.0 36.7 5.6 13.3 16. 18.8 38.8 25.5 21792 86.4 44.6 58.7 43.2 6.7 16.7 17.7 26.3 55.6 40.5 21794 82.2 41.3 53.1 44.1 8.2 16.3 16.8 23.6 53.0 36.1 21935 69.5 44.0 41.1 36.0 5.5 13.9 15.5 19.4 39.3 25.7 21937* 93.0 53.5 62.0 48.0 4.3 19.6 20.8 27.5 21961 82.4 40.0 56.5 46.5 7.4 17.1 19.2 24.5 53.2 40.1 22001 78.1 40.1 49.5 40.3 5.6 15.0 18.0 22.0 49.1 33.3 Females 21786 81.4 42.9 52.3 42.7 7.9 16.0 18.0 23.4 50.5 36.2 21776 75.2 43.2 45.7 37.5 5.93 13.8 16.8 21.5 46.1 29.2 21777 76.2 42.9 46.1 37.4 5.4 15.0 16.7 20.1 45.6 30.3 21780 85.1 41.9 53.1 44.3 7.1 16.6 18.7 22.6 53.7 38.3 21778 78.8 40.9 49.6 41.3 7.1 16.3 18.0 21.2 49.5 39.6 21789 79.2 41.8 48.9 41.2 6.4 13.9 17.0 21.9 50.4 33.4 21782 82.5 43.2 54.3 45.3 7.7 17.2 18.1 23.9 52.2 38.4 21788 80.7 41.3 51.1 43.4 6.9 16.9 18.4 23.3 53.7 36.0 21790 76.2 43.0 47.3 39.3 6.9 14.6 16.2 21.2 46.1 30.2 21787 82.1 43.1 51.8 44.5 7.6 17.2 17.8 23.6 51.4 34.7 21793 69.5 42.3 35.1 35.1 5.4 13.8 15.8 18.1 34.4 23.0 21934 79.4 41.1 52.4 41.0 6.1 16.0 18.1 23.1 49.2 27.5 21938 71.5 39.8 49.7 38.3 4.5 15.8 18.0 20.5 44.4 34.2 21960 80.8 42.3 50.5 40.1 6.25 15.5 17.5 21.6 50.2 35.8 23039 68.3 40.1 38.8 33.0 4.2 14.1 15.8 16.7 35.4 19.5* Correct skull for specimen? Collected by S.F. Ferrari.

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155 Map 18. Location map for Jamari sakis. Brazil 1. UHE Samuel, Schneider collection site 2. Rio Jamari, Usina Hidroeltrica Samuel 3. Schneider and Junior collection sites (approx.) 4. Braga and Souza Jnior collection sites (approx.)

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156 Figure 103. Key specimen MPEG No. 21961, adult male, skull Figure 102. Key specimen MPEG No. 21961, adult male, skin.

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Neotropical Primates 21(1), July 2014 157 Figure 105. Key specimen MPEG No. 21785, adult male, skull. Figure 104. Key specimen MPEG No. 21785, adult male, skin. Figure 106. Key specimen adult female MPEG No. 21934, skin. Figure 107. Key specimen adult female MPEG No. 21787, skin.

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Neotropical Primates 21(1), July 2014 158 Introductione following is a reinterpretation of the paintings commis sioned by Philip Hershkovitz with artist Zorica Dabich for his unnished volume Living New World Monkeys, Volume 2 (Hershkovitz died before it was completed). All images used are by Zorica Dabich and Philip Hershkovitz, through the courtesy of Field Museum of Natural History. ese images were published in Hershkovitz (1987) as Figure 17. e caption for that image read: Fig. 17. Pithecia: Males of all species (and subspe cies of P. irrorata) and female of P. pithecia pithecia, all to same scale; an arrangement of photographs of individual paintings in color by Zorica Dabich (from Hershkovitz, Living New World Monkeys, Volume 2 [in preparation]). Dierences between subspecies of P. monachus and P. pithecia are not apparent in the black and white gures. e gure shows the paintings in order of: P. monachus, P. i. irrorata, P. i. vanzolinii, P. albicans, P. aequatorialis, P. pithecia (female), and P. pithecia (male). All of the corresponding art for comparison in this appen dix is by Stephen Nash. A III. U F 17 P. Hrfn 1987: Tr Tt Sr Ab Sf, G PITHECIA (C, Ptr): A Pt R Cb R Wr r Db N Sb N Sb, AM. J. PRIMATOL 12:387Painting A: P. monachus is image depicts an amalgam of saki traits that lead to some of the confusion that was P. monachus throughout the Amazon (cf. e Monachus Mess in P. monachus Discussion). e species this painting most closely resembles is the newly described Pithecia isabela as some members of the spe cies can have a faded white wash forehead instead of the more distinct white eye spots and have a minimal amount of white stippling throughout the body pelage (cf. P. isabela). Alternatively, this image is a catchall of generic female char acteristics, but it is unclear for which species. It was this painting, that turned P. monachus into a clumping of everything, male and female, that might t this ambiguous image, especially since it was published in black and white. What Hershkovitz did not realize is that the type for monachus, a juvenile of indeterminate sex with a white band around its face, would not grow up and retain that color pattern, but would transition, like many of the species do, into a form that looks completely dierent as an adult. As a comparison, at left are P. hirsuta, P. monachus, and P. isabela, the likely candidates that contributed to this blended image.

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Neotropical Primates 21(1), July 2014 159 Painting B: P. irrorata is image of P. irrorata is another amalgam because Hershkovitz did not know there was more than one baldfaced saki species when he published in 1987. is image by itself looks most similar to a subadult P. mittermeieri, as does the photo in Hershkovitz 1987 in Figure 25 (cf. Table 2, Introduction). As a comparison, below are illustrations of P. irrorata and P. mittermeieri. is juvenile male P. mittermeieri appears to be the most similar to the painting from Hershkovitz (1987). Photo by L.C. Marigo (Figure 85a, P. mittermeieri).

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Neotropical Primates 21(1), July 2014 160 Painting C: P. vanzolinii is painting of a P. vanzolinii male was based on museum material and fairly well represents the species. It is unlikely that Hershkovitz ever saw any of these animals alive, which is the same challenge for this taxonomy. e Nash version of male and female is pictured at the right. Painting D: P. albicans is is an interesting depiction of P. albicans in that the males and females are very similar in pelage, but it is the females that have the white facial hair. us, this painting is actually of a female P. albicans. Male P. albicans have a mostly plain face with creamcolored eyebrows, as the illustrations at right demonstrate. Note on the close up of the male face, the cream-colored hair appears only on the forehead of the male.

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161 Painting E: P. aequatorialis is lovely painting, the original of which hangs in the base ment hallways of e Field Museum, is a nearly accurate de piction of Hershkovitzs P aequatorialis. e only thing miss ing is the distinct orange ruf on the male. At the right for comparison, is Nashs version of the male and female.

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Neotropical Primates 21(1), July 2014 162 Painting F: P. pithecia male. Painting G: P. pithecia female The most recognizable and most collected of all of the Pithecia species, P. pithecia is perfectly well represented in these two accurate paintings. The nearly identical Nash version is at right.

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163 Philip Hershkovitz, Emeritus Curator of Mammals in his laboratory at the Department of Mammalogy in the Field Museum of Natu ral History in Chicago (1.987). Photo by Stephen Nash.

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Neotropical PrimatesA Journal of the Neotropical Section of the IUCN/SSC Primate Specialist Group Conservation International 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USAISSN 1413-4703 Abbreviation: Neotrop. Primates Editors Erwin Palacios, Conservacin Internacional Colombia, Bogot DC, Colombia Liliana Corts Ortiz, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA Jlio Csar Bicca-Marques, Pontifcia Universidade Catlica do Rio Grande do Sul, Porto Alegre, Brasil Eckhard Heymann, Deutsches Primatenzentrum, Gttingen, Germany Jessica Lynch Alfaro, Institute for Society and Genetics, University of California-Los Angeles, Los Angeles, CA, USA Anita Stone, Department of Biology, Eastern Michigan University, Ypsilanti, MI, USA News and Books Reviews Brenda Solrzano, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Ernesto Rodrguez-Luna, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Founding Editors Anthony B. Rylands, Center for Applied Biodiversity Science Conservation International, Arlington VA, USA Ernesto Rodrguez-Luna, Instituto de Neuroetologa, Universidad Veracruzana, Xalapa, Mxico Editorial Board Bruna Bezerra, University of Louisville, Louisville, KY, USA Hannah M. Buchanan-Smith, University of Stirling, Stirling, Scotland, UK Adelmar F. Coimbra-Filho, Academia Brasileira de Cincias, Rio de Janeiro, Brazil Carolyn M. Crockett, Regional Primate Research Center, University of Washington, Seattle, WA, USA Stephen F. Ferrari, Universidade Federal do Sergipe, Aracaj, Brazil Russell A. Mittermeier, Conservation International, Arlington, VA, USA Marta D. Mudry, Universidad de Buenos Aires, Argentina Anthony Rylands, Conservation International, Arlington, VA, USA Horcio Schneider, Universidade Federal do Par, Campus Universitrio de Bragana, Brazil Karen B. Strier, University of Wisconsin, Madison, WI, USA Maria Emlia Yamamoto, Universidade Federal do Rio Grande do Norte, Natal, Brazil Primate Specialist Group Chairman, Russell A. Mittermeier Deputy Chair, Anthony B. Rylands Vice Chair, Special Section on Great Apes, Liz Williamson Vice Chair, Special Section on Small Apes, Benjamin M. Rawson Regional Vice ChairsNeotropics Mesoamerica, Liliana Corts-Ortiz Andean Countries, Erwin Palacios and Eckhard W. Heymann Brazil and the Guianas, M. Ceclia M. Kierul, Fabiano R. de Melo and Mauricio Talebi Regional Vice Chairs Africa W. Scott McGraw, Janette Wallis and David N.M. Mbora Regional Vice Chairs Madagascar Christoph Schwitzer and Jonah Ratsimbazafy Regional Vice Chairs Asia China, Long Yongcheng Southeast Asia, Jatna Supriatna, Christian Roos, Ramesh Boonratana, and Benjamin M. Rawson South Asia, Sally Walker and Sanjay Molur Layout: Kim Meek, Washington, DC IUCN/SSC Primate Specialist Group logo courtesy of Stephen D. Nash, 2002. Front cover: Adult male Pithecia hirsuta, Capar Biological Station, Colombia. Photo by R.A. Mittermeier. is issue of Neotropical Primates was kindly sponsored by the Margot Marsh Biodiversity Foundation, 432 Walker Road, Great Falls, Virginia 22066, USA, and the Los Angeles Zoo, Director John R. Lewis, 5333 Zoo Drive, Los Angeles, California 90027, USA. Printed in Bogot DC, Colombia on July 25, 2014. NP 20_supplement_COVER_k.indd 2 7/25/14 10:33 PM Scope e journal/newsletter aims to provide a basis for conservation information relating to the primates of the Neotropics. We welcome texts on any aspect of primate conservation, including articles, thesis abstracts, news items, recent events, recent publications, primatological society information and suchlike. Submissions Please send all English and Spanish contributions to: Erwin Palacios, Conservacin Internacional Colombia, Carrera 13 # 71-41 Bogot D.C., Colombia, Tel: (571) 345-2852/54, Fax: (571) 3452852/54, e-mail: , and all Portuguese contributions to: Jlio Csar Bicca-Marques, Departamento de Biodiversidade e Ecologia, Pontifcia Universidade Catlica do Rio Grande do Sul, Av. Ipiranga, 6681 Prdio 12A, Porto Alegre, RS 90619-900, Brasil, Tel: (55) (51) 3320-3545 ext. 4742, Fax: (55) (51) 3320-3612, e-mail: . Contributions Manuscripts may be in English, Spanish or Portuguese, and should be double-spaced and accompanied by the text on CD for PC compatible text-editors (MS-Word, WordPerfect, Excel, and Access), and/or e-mailed to (English, Spanish) or (Portuguese). Hard copies should be supplied for all gures (illustrations and maps) and tables. e full name and address for each author should be included. Please avoid abbreviations and acronyms without the name in full. Authors whose rst language is not English should please have their English manuscripts carefully reviewed by a native English speaker. Articles. Each issue of Neotropical Primates will include up to three full articles, limited to the following topics: Taxonomy, Systematics, Genetics (when relevant for systematics and conservation), Biogeography, Ecology and Conservation. Text for full articles should be typewritten, double-spaced with no less than 12 cpi font (preferably Times New Roman) and 3-cm margins throughout, and should not exceed 25 pages in length (including references). Please include an abstract in the same language as the rest of the text (English, Spanish or Portuguese) and (optional) one in Portuguese or Spanish (if the text is written in English) or English (if the text is written in Spanish or Portuguese). Tables and illustrations should be limited to six, except in cases where they are fundamental for the text (as in species descriptions, for example). Full articles will be sent out for peer-review. For articles that include protein or nucleic acid sequences, authors must deposit data in a publicly available database such as GenBank/EMBL/ DNA Data Bank of Japan, Brookhaven, or Swiss-Prot, and provide an accession number for inclusion in the published paper. Short articles. ese manuscripts are usually reviewed only by the editors. A broader range of topics is encouraged, including such as behavioral research, in the interests of informing on general research activities that contribute to our understanding of platyrrhines. We encourage reports on projects and conservation and research programs (who, what, where, when, why, etc.) and most particularly information on geographical distributions, locality records, and protected areas and the primates that occur in them. Text should be typewritten, doublespaced with no less than 12 cpi (preferably Times New Roman) font and 3-cm margins throughout, and should not exceed 12 pages in length (including references). Figures and maps. Articles may include small black-andwhite photographs, high-quality gures, and high-quality maps. (Resolution: 300 dpi. Column widths: one-column = 8-cm wide; two-columns = 17-cm wide). Please keep these to a minimum. We stress the importance of providing maps that are publishable. Tables. Tables should be double-spaced, using font size 10, and prepared with MS Word. Each table should have a brief title. News items. Please send us information on projects, eld sites, courses, esis or Dissertations recently defended, recent publications, awards, events, activities of Primate Societies, etc. References. Examples of house style may be found throughout this journal. In-text citations should be rst ordered chronologically and then in alphabetical order. For example, (Fritz, 1970; Albert, 1980, 2004; Oates, 1981; Roberts, 2000; Smith, 2000; Albert et al., 2001) In the list of references, the title of the article, name of the journal, and editorial should be written in the same language as they were published. All conjunctions and prepositions (i.e., and, In) should be written in the same language as rest of the manuscript (i.e., y or e, En or Em). is also applies for other text in references (such as PhD thesis, accessed see below). Please refer to these examples when listing references: Journal article Stallings, J. D. and Mittermeier, R. A. 1983. e black-tailed marmoset ( Callithrix argentata melanura) recorded from Paraguay. Am. J. Primatol. 4: 159. Chapter in book Brockelman, W. Y. and Ali, R. 1987. Methods of surveying and sampling forest primate populations. In: Primate Conservation in the Tropical Rain Forest, C. W. Marsh and R. A. Mittermeier (eds.), pp.23. Alan R. Liss, New York. Book Napier, P. H. 1976. Catalogue of Primates in the British Museum (Natural History). Part 1: Families Callitrichidae and Cebidae. British Museum (Natural History), London. esis/Dissertation Wallace, R. B. 1998. e behavioural ecology of black spider monkeys in north-eastern Bolivia. Doctoral thesis, University of Liverpool, Liverpool, UK. Report Muckenhirn, N. A., Mortensen, B. K., Vessey, S., Fraser, C. E. O. and Singh, B. 1975. Report on a primate survey in Guyana. Unpublished report, Pan American Health Organization, Washington, DC. Website UNESCO. 2005. UNESCO Man and the Biosphere Programme. United Nations Educational, Scientic, and Cultural Organisation (UNESCO), Paris. Website: http://www.unesco.org/mab/index.htm. Accessed 25 April 2005. (Acessada em 25 de abril de 2005 and Consultado el 25 de abril de 2005 for articles in Portuguese and Spanish respectively). For references in Portuguese and Spanish: and changes to e and y for articles in Portuguese and Spanish respectively. In changes to Em and En for articles in Portuguese and Spanish respectively. Doctoral thesis changes to Tese de Doutoramento and Tesis de Doctorado for articles in Portuguese and Spanish respectively. MSc esis changes to Dissertao de Mestrado and Tesis de Maestra for articles in Portuguese and Spanish respectively. Unpublished report changes to Relatrio Tcnico and Reporte no publicado for articles in Portuguese and Spanish respectively. Notes to Contributors NP 20_supplement_COVER_k.indd 3 7/25/14 10:33 PM

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NEOTROPICAL PRIMATES rfnnftbnfbrfn bfntnbbft n n bfbb bbffb bbn n nbr bffnnnbnn nfnbnbffnnbnntfbf Neotropical Primates A Journal and Newsletter of the IUCN/SSC Primate Specialist Group Vol. 21(1), July 2014ContentsA Taxonomic Revision of the Saki Monkeys, Pithecia Desmarest, 1804Laura K. Marsh .................................................................................................................................................................1 NEOTROPICAL PRIMATES rfntbb