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Home , Kipunji, Sanje mangabey

Kitulo Plateau and adjacl'Tlt Livingstone Forest The Highland as the Kitulo National Park (412 km1• O9°OO'S to 09' 16'S and JJ'4J'E to J4'OJ"E) (2). Lophocebus : A New Mount Rungwe Forest Reserye (150 km2, 1)'J'OJ'S 10 09' 12' Sand 33'35'1' to 33'45'E) of African Monkey supports montane and upper montane fmest, bamboo. and plateau grassland. The montane Trevor Jones," Carolyn L Ehardt,2 Thomas M. Butynski,' forests of Mount Rungwe and Livingstone Tim R. B. Davenport,4 Noah E. Mpunga: Sophy J. Machaga: (Rungwe-Livingstonc) arc joined only by a narrow corridor of degraded forest. Until now, Daniela W. De Luca4 eight species of were known from the A distinct species of mangabey was independently found at two sites 370 Southern Highlands, including a probable new kilometers apart in southern Tanzania (Mount Rungwe and Livingstone in the species of dwarf galago, Galagoides sp. (3). Southern Highlands and Ndundulu in the ). This new species During interviews in January 2003 in vil- is described here and given the name Mhighlandmangabey" Lophocebus kipunji sp. lages around Mount Rungwc. we heard rumors nov. We place this monkey in Lophocebus. because it possesses noncontrasting aboul a shy and alypical monkey known as black eyeUds and is arboreal L kipunji is distinguished from other mangabeys by Kipunji (kip-oon-jee). The local Wanyakyusa the color of its pelage; long. upright crest; off-white tail and ventrum; and loud have a strong oral trddition based on both real call This find has implications for primate evolution, African biogeography, and and mythical forest . and validation of forest conservation. these rumors was protracted. We first ob- served an unusual primate during biodiversity 1bc most recently discovered .species of monkey Southern Highlands population. The surveys on f\..'ount Rungwe in May 2003. but in Afiica was the sun-tailed monkey, Cercopithe- Southern Highlands of southwest Tanzania cuS'solatus, fOlmd in Gabon in 1984 (I). 'Jere, (Fig. 1) rise to 2961 m above sea level (asl) lUdzungwa Mountains National Park, Box 99. Mang'ula. we report on the discovery of a new species of and comprise mountain ranges capped by Tanzania. 'Department of Anthropology, University mangabey in Tanzania. 11lis discover)' wa.<;made forest-grassland mosaic. The Highlands re- 01 Georgia. Athens, GA 30602-1619, USA. 3ConSf'f- almost simultaneously by independenl field- ceive rainfall via convectional uplil1 from vat ion International, Post Office Box 68200. City Square 00200, Nairobi, Kenya. "wildlife Conservation workers on different mOWltainmngcs in southt.-m Lake Nyas..1.of up to 2900 mill a year, the Society, Southern Highlands Conservation Pro- Tanzania. We relate the circumstanct.'S of dis- highest in Tan/-,mia. gramme, Post Office Box 147S. Mbeya, Tanzania. covery in the two sites, describe and name the Within the Southern Highlands. the Tanza- .To whom correspondence should be addresspd. new species, and discuss its . nian government is presently gazetting the E-mail: ~ G 6 • C~ \ce.-•..Jl.+ .0 Uo..'j '" .e.<.~ C1 'Z...... -rc.\ '('" . ~ ,",-", r).4 ~ u.-t.s~

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>r ~c A '" ••••• B M' ••••• - 0 MBEYA 0

If!:INI3A 0 ,-- - ~

QT_ • • D •••.•••••) .( OF_A_ ~ -' ~ 0 •••••••• f ••••• - '''''''''' .~ •••• of - L.~ ,- , N = " "'~ - • " ".. - w~e >r • - - - -(TA80'Oot) Fig. 1. Maps of Tanzania's Udzungwa Mountains (A) and Southern Highlands (8) showing the known range of the highland mangabey Lophocebus kipunji.

because of the terrain, thick secondary for. cst, and the 's cryptic nature, sightings were infrequent and poor. It was not until De~ ccmber 2003, during work in the contiguous Livingstone Forest, that the monkey was clear- ly observed and recognized as a new species of rnangabcy. Ndundulu population. The Udzungwa MOWltain, (Udzungwa." 10,000 lan', 07°40'S to 08°40'S and 35° IO'E to 36°50'E) lie 350 Ian 10 the northeast of Rungwe-Livingstonc (4, 5). Supporting cin:a (ca.) 1017 lan' of fragmented forest (6), the Udzungwas receive a maximum annual rainfall of roughly 2200 mm and were previously thought to hold 10 primate spccil."S (7), including lhe endemic , Ccrcocelms sanjei, discovered in 1979 (8). Two populations of the Sanjc m.mgabey are known from the UdzlUl&'\'as (7, 9). During Fig. 3. Paratype: adult highland mangabey visits from 1991 to 2000, ornithologists work- Lophocebus kipunji, Ndundulu Forest Reserve. ing in the Ndundulu Forest R~er\'e (Fig. I) Tanzania. [Photograph by T. Jones] reported a third population of the Sanje mangabcy (10). Subsequent surveys failed to tail off-white. Crown with very long. broad, conlinn the presence of this species in crect crest of hair. Eyelid ••black, not contrast- Ndundulu (7, 9, J J) and led to our intensified Fig. 2. Holotype: adult male highland manga- ing with color of face. Adults emit a distinc- surveys in July and September 2004. During bey Lophocebus kipunji in the type locality. tive, loud, low-pitched "honk-bark" (Fig. 4). these surveys, Sanje mangabeys were not Rungwe-Uvingstone. Tanzania. [Photograph by Alboreal. FUWldonly at high altitudes (1300 m encountered or heard. Ilowever, on 7 JulY T.R.B.Davenport] up to 2450 m asl) and low-temperature tol- 2()()4, the new species of mangabcy was dis- erant; temperatures in Rungwe-Livingstone covered. It now seems certain that the orni- live individual should be collected at this drop to at least -3°e. thologists had misidentified the new species time to serve as the holotype. The Rungwe- Description. A primarily brown. mediwn- of mangabey as the Sanje mangabey. Livingstone population is designated the sized, long-tailed, arboreal monkey. Muzzle The researchers working on each of these source population for physical specimens in elongated. Facial skin. including eyelids, two new populations of mangabeys did not support of the holotype. black. Suborbital fossae "tear line" pro- become aware that a second population was Paratype, Adult in photograph (Fig. 3). nounced. Eyes broM!o Pelage light to rufous known until October 2004. Sex not known. Photograph taken in Ndundulu brown except as follows: center of vcntrum Lophocebus kipunji Ehardt, Butynski, Forest Reserve (07°48'45"S 36°31'05"E), and distal half of tail, white to off-white; hands Jones, and Davenport sp. nov. Udzungwa Mountains, Tanzania. and feet, black; lower forelimbs, dark brown to Holotype. Adult male in photograph Type locality. Rungwe-Livingstone black. Cheek whiskers long. Crown with vel)' (Fig. 2). Photograph taken in the type (09°07'S to 09°II'S and 33°40'E to 33°55'E), long. broad, stilT, upright crest of hair. Shoul~ locality at 9°07'S 33°44'E (12). The number Southern Highlands, Tanzania. der cape prescnt in some individuals. although of individuals in each of the two populations Diagnosis. Pelage of dorsum light to medi- there is variation in length and color. White of of this species is undoubtedly very small; no wn brown., center ofventrum and distal half of vcntrum sharply offset from brown in at least

1162 20 MAY 200S VOL 308 SCIENCE www.sciencemag.org REPORTS kHz "'N''" 1...kipunji arc known from only 3 km2 of 5 Ndundulu Forest. Previous rl.'Search in the Bark. urils area (7, 11,22, 23) and lack of independent 4 ~Barl<.trvtsil i I I ~ i knowledge of the species among local villag- 3 M~ ers indicate that this monkey is ab~nt from 2 large parts of this forest. We e:-.1imate the geographic range for this populalion to be 3 to 1 ~. 50 km2• Only three groups have so far been located in Ndundulu. and it seems unlikely 1V 0 5 5 thai this population exceeds 500 animals. Fig. 4. Sonogram of three honk-bark. calls given by adult L. kipunji from the type locality. with two Once lhc degree of threat status is fonnally series of barks. Honk-barks. which appear to playa role in intergroup spacing. are given singly or in assessed for the IUCN Red List of Thrmtened series and may. as in this example. be interspersed with barks. Honk-barks are short discrete phrases, composed of a variable number of units. The first honk.bark shown here has six units. Animals, we expect that L. kipun)i will he clao;- Honk..barks are frequency modulated with formants or emphasized harmonics. In the honk.bark. sifir...'das a critically (24). the first harmonic or "fundamental frequency" is that part of the caU with the lowest frequency The presence of a third threatened species and often the greatest amplitude. and is thus the farthest traveling element. The fundamental of monkey makes the Udzungwas arguably frequency of the honk-bark is 0.28 kHz (n = 3). the frequency range is 0.180 to 0.304 kHz. and the the most important single site in Africa for the mean unit interval is 0.076 s (n = 3). This call is thus qualitatively and quantifiably different from conservation of primate biodiversity. The the type I and type II calls of other studied mangabey species (20. 21). [Call recorded by N. E. Mpunga. Sonogram prepared and interpreted by A. Perkin.] presence of L. kipun)i in both sites also sup- ports the hypothesis that the Southern High- lands are zoologically more aligned to the some Ndundulu animals. Ventrum lighter than and are semiterrcstrial. LophocehrL'l spp. are Eastern Arc Mountains than has usually been rest of body in Rungwe-Livingstone animals most closely related to haboons, Papio spp., thought (25). but not sharply on:••et. Small patch of ofT-white and the . TheropitheCIL\' ge!ada; have The threats to 1... J."ipunjiarc considerable. hair between neck and stcrnum in Rungwe- black eyelids that do not contmst with the TIle Rungwe-l.ivingstonc forests are severely Livingstone animals. Individual hairs appear to color of the face; and are arboreal (14, 16-18). dcgmdl'O. Logging, charcoal making. poach- be straight and devoid of bands or speckling. We place this new species in the genus ing, and unmanagl.-d Tt.'SOurceextraction are Tail pelage smooth. not shaggy or lax. Tail car- Lophoce"'L~ on the basis of the combination common. The narrow forest corridors linking ried loosely and parallel to or below plane of the of its noncontrasting black eyelids and arbore- Mounl RWlgwe to Li\oingSlone ;:mdjoining the back, curving downward to fl'et level during ality. 1... kipunji is isolated from the nearest northern and southern sections of Livingstone locomotion. Tail not held strongly arched over other species of Lophocebus (1...albigena) by arc almost completely encroached upon. With- back when standing, nor held vertically, except ahout gOO km. out immediate conservation intervention, these a'i a semi-prehensile support. No sexual dimor- The other two species of Lophocebus (15) forests will be lragmented. resulting in three phism in color of adult pelage. Ischial callosities differ from L. kipunji as follows: isolated suhjXlpulalions of L. kipunji; indeed, pink: fused in the male, unfiJscd in the female. Gray-checked mangabey L. albigena: the ca.'>1r...mmost:min1alsmay already be isolated. Genital swelling in oestrus females. Body and tail blackish-brown. Check whisk- Gazcllemcnt of Kitulo National Park should help Measurements. None available. Head- ers gray. Crown with short tuft (or tufts) of protect the !lomalkr Livin~~1onc Forest popula- body length of adult males in the Rungwe- hair (14). Frequently hold'i tail vertical or tion. However, sv.ill and effective action is Livingstone population estimated at 85 to 90 em. arched over back. "Whoop-gobble" loud call. impl.'T3tive.bulb in the new park and especially Tail about equal in length to h""d and lxxly. !\!1ainly occurs below 1600 m (/3), except \\ithin Mount Rungwe Forest Rest....rve. Adult male body weight estimated at 10to 16kg. Nywlgwe. Rwanda, and Kahuzi.Biega. Dem- Although the submontanc forest ofNdundulu Etymology. The specific name acknowl- ocratic Republic of Congo, where it occurs up is in excellent condition and no signs of recent edges the Kinyakyusa name for this monkey to ca. 2350 m (N). hunting have been observed in the area from in Rungwe-Livingstone. Black mangabey L. aterr;mus: Entire which L. bpunji is kno'om. this monkey is pres-- Distribution. Kno,,",llto occur in ca. 70 km2 body black. Crown with tall. thin, central ent in low numbers. If lhis ~lllall population is of Rungy,e.Li"ingstone in the Southern High- luft of hair. Check whiskers thick and gray to he protected in perpetuity, the Udzungwa lands, Tanzania, from 1750 up 10 2450 m asl (14). Whoop-gobble loud call. Not known to Mountains National Park needs to be extended (possibly up to 2600 m asl), and from ca. 3 occur above 450 mas!. to include the Ndundulu Forest. km2 of Ndundulu Forest Reserve, Tanzania. All mangabey species ~tudied to date from 1300 up to 1750 m asl. pOSSL'SSa loud call, emitted by adult males to References and Notes Habitat. From pristine submontane forest coordinate intergroup spacing (20. 2 J). The 1. M. J. S. Harrison.). lool. Z15. 561 (1988). in Ndundulu to degraded montane and upper characteristic whoop-gobble loud call of other 2. T. R. B. Davenport. Oryx 36. 224 (2002). montane forest in Rungwe-Livingslone. Lophocelms species has not been detected in 1... 3. A. Perkin. personal communication. Mangabeys arc medium.sized monkeys kipulI)i. Adult males in RWlgwe-Livingstone 4. N. Myers. R. A. Mittermeier. c. G. Mittermeier. G. A. B. da Fonseca. J. Kent, NawfP 403. 853 (2000). confined to the forests of equatorial Africa do give a di~tinclive honk-bark (Fig. 4). which 5 T. M. Brooks et al .• ComeN. Bioi. 16.909 (2002). (/3). Taxonomically controversial, there arc is most evident when conspecific groups 6. N. D. Burgess. J. FjeldsA, R. Botterwpg. j. East AIr. currently from five to nine species (and from meet. Nat. Hist. B7. 37 (1998). 7. C. L Ehardt. AIr. ., 15 (ZOO1). five to II subspecies) recognized in two In RWlgy.o'e-Livingstone,we have identified 8, K. M. Homewood. W. A. Rodgers, Int. j. Pr;matol. Z. gl."J1em(Loph(}(:ehus and Cercocehus) (/4. 15). J() groups of L. kipun)i: five in the southern 47 (1981). The mangabcys are diphyletic. as revealed by seclion of MOWlt Rungwe. four in the north. 9. C. L Ehardt. T. Jones, T. M. Butynski. Int. j. Primatof. Z6. 557 (2005). molecular studies and supported by anatomi- em section of Livingstone, and one in the 10. L. Dinesen. T.lehmberg. M. C. Rahner. J. Fjeldsa. Bioi. cal and behavioral characteristics. CercoceblL~ southern section of Livingstone 20 km to the ConseN. 99,223 (2001). spp. are most closely related to and east (Fig. I). We estimate Ihe total population 11. C. L Ehardt, T. T. Struhsaker. T. M. Sutynski. 'Conser. vation of the Endangered Primates of the Udzungwa (J.J{lndrj/hL~ drills spp.), have pink or white of L. kipunji in Rlrn£'\'e-Livingstone to be Mountains, Tanzania: Surveys. Habitat ASSt'!isment, and eyelids that contrast with the color of the face, 250 to 500 animals. Long-Term Monitoring" (Margot MaM Biodiv~ity

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Foundation, Great FaUs,VA, and Conservation Interna- diversity Conservation Project. Matumizi Ende-levu ya thank SHCP staff, R. Ngindale, G. Patterson, Rungwe tionaL Washington, DC, 1999). Misitu ya Asili, lringa, Tanzania, 20(1). District Council P. Chibwayl!', Tanzania National 12. A. Wakeham-Dawson, S. Morris. P. Tubbs, Bult. Zool. 23. A R. Marshall, J. E. Topp-jorgensen. H. Brink, E. Fanning. Parks (TANAPA), W. Mwakilema, Z. Tweve, J. Maher, Nomencl. 59, 18 (2002). Int. J. Primatol. 26, 127 (ZOO5). and B. Stanley. Fil'ldwofk in Ndundulu (SMRP) was 13. 1. Kingdon, The Kingdon Field Guide to African 24. fUCN Red Ust Categories arnJ Criteria: Version 3. J supported by WCS, Primate Society of Great Britain, (Academic Press, london, 1997). (IUCN-The Wortd Conservation Union Species Survival Critical Ecosystem Partnership Fund, Conservation 14_ C. P. Groves, Primate (Smithsonian, Commission, IUCN, Gland, Switzerland, 20(1). International, Primate Action Fund, the Margot Washington, DC. 2001). 25. T. R. B. Davenport, Soc. ConseN. Bioi. Abst. 048, 20 Marsh Biodiversity Foundation, the University of 15. P. Grubb et .11.,Int j. Primatol. 24. 1301 (2003). (2004). Georgia Research Foundation, and Primate Conser- 16. C. P, Groves. Primates 19, 1 (1978). 26. CLE. (Sanje Mangabey Research Project. SMRP) and vation, Incorporated. We thank A. Alchard, R. Laiuer, 17. E. l Hams, T. R.Di'SOteU,ftfol. BioI, EvoI.. 15,892 (1998). T.R.B.D. (Southern Highlands Conservation Pro- A. Mndeml!', Tanzanian Wildlife Research Institute, 1S. J. G. Fleagle. S. W, McGraw. Proc. Nat/. Acad. Sci. gramme, SHCP) directed the projects that produced Tanaunia Commission for SCience and Technology, U.S.A. 96, 1157 (1999). the discoveries. T.J. located and observed the TANAPA, Iringa District Counci\, J. Massao, I. Chahe, 19 R. J. Dowsett. F. Dowsett-lemaire. in Survey of the Ndundulu population. T.J~ CLE., and T.M.B. contri- }. Mudanga, Q. Lukl!', and the two anonymous re. Fauna and Flora of Nyungwe Forest Rwdnda. R. J. buted data for thl' Ndundulu population; T.R.B.D., viewers of the manusaipt. Particular thanks to L Dowsett, Ed. (Tauraco. Ely, UK, 1990), pp. 111-121. N.E.M., S.l.M., and D.W.D. located, observed, and Dinesen, T. lehmberg. and A Marshall for locational 20. P. M. Waser, in Primate Communication, T. Snowdon. contributed data for the Rungwe-Uvingstone popu- information on Ndundulu; A. Perkin for pr~aring thl!' C. H. Brown, M. R. Peterson, Edt. (Cambridge Univ. lation; T.J., CLE., T.M.B., and T.R.B.D. wrote thl' paper sonogram; and A Polaszek for expert advice on Press, Cambridge. 1982). pp. 117-144. with editorial input from N.E.M., S.J.M., and D.w.D. behalf of thl!' International Cornmissioo 00 Zoolog- 21. F. Range, J. Fischer, Ethology 110. 301 (2004). Fieldwork. in Rungwe-livingstonl!' was funded by the ical Nomenclatune. 22. Ie:. z. Doody, K. M. Howell, E. Fanning. Eds~ .West Wildlife Consl!'(\'ation Society (WCS) through the Kilombero Scarp Forest Reserve zoological report- SHCP with the generosity of E. McBean, G. Fink, 27 December 2004; accepted 3 March ZOOS (Udzungwa Mountains Forest Management and Bio- Nikon Incorporated, and an allOnymous donor. We 10.1 126Jscience.1 109191

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