FIELDLAN Zoology NEW SERIES, NO. 113 Systematic Review of the Barbary Macaque, Macaca sylvanus (Linnaeus, 1758) Jack Fooden October 26, 2007 Publication 1547 . PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Bs ina ; Rite OL F ») me mete cb St Y ai Foley las RI ee ALS Mi, ee A ae Wad C » fd zt Rie Ma OS FIELDIANA Zoology NEW SERIES, NO. 113 Systematic Review of the Barbary Macaque, Macaca sylvanus (Linnaeus, 1758) Jack Fooden Division of Mammals Department of Zoology Field Museum of Natural History 1400 South Lake Shore Drive Chicago, Illinois 60605-2496 U.S.A. Accepted July 20, 2007 Published October 26, 2007 Publication 1547 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2007 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents ADIT SYNC tees scp ay Biecoate se AC cn a icine tire ea cad cs es erat Ol tar Ot wh ee han Sa rare eRe aS idee i Se dee a ] ENTRODUCIIONSEETE Mente recreate eer rere Meme tee chore mate k ene: eens tiie ites chitis ge fie a ace tts Case Renae eter 1 EATER TALSPAN DE IVER EOS oo. ncltora tare tete cotorererel chert nerions verre er tele ee en queers OP nile lade oa eGR) Gulewe io emearente l ea NASER ASTI Be NE SITER TENA UMN otc esse Seog adetr eter hcescraas te han rah at aihcg taba hans irate tata ha aie Nia ieus' ahs" Sabra andl ol ERD 2 iA eE OPULATTONSESTINMAMESS Aer tecttatcoahet cucict ara stun Orch s sei amet coseus 4) Midis ei MPSC EEA Ses ecu sere eteene 2 Op OBES 0 ta flighty BUEN a EI a OT Rng AES Se en a MERE Tree ae EY 5 ESGIERNAT: MIEASUREMENTIS{ANDMEROPORTIONS © 40 5.¢ 02s slaue le he oe soe) Sache oie eyepe cie eG) eee ce ey ee au an auc 4 A DATE ERTEBRAE AC Wena cuter beet tayhe seer aalt sales ores \Gurabeee a eth cAI suse dats ie asa ot) oa rtati, eleoee enon earns 6 Sena RAE OT AUTRE Rte Se 1 Pant ee ntl Mae ve let eligte caudate aw duh Co Gree Ge wetin eee a ea ia pea ae pe 6 MOUECULARGBIOLOGYAAND I GENETICS S42 Un elke. ates tard een ene he hte ere Chee, De ete eA OMe IN pie an iene te 8 “Le gy Tra 2 nome ete a ata Spe ce We ein Serra a OP COENEN OP EE RIC od de aie as aba te fea nt We 8 Mee te eh te rere ete he See Teta ada ety he wtatattn Catan aot al alee etm a ced See eee he PSI AUTIUG RUAN Ee TSU) Ravocpee Revre gue St he teas WN, ga oe A a et IS is ed maa 2 Moho pas astantap tay Ben nme yemenene, Me creates 13 [Sie vhs oe ee eB Cer or Saray ee pee eat etc 7 perme PR Pa 3 PRESSE Hee EHR, PER Mk cmt ACh 13 Piers A COCCI ic! Pye atk Te ee FEB Loui Mitac Taped ta da lagtue ene ee shay Pu cnaus athe te) pe center 14 remue site anid OID astON poe er Cons te ete rete oats) ko niece a aus Sate a nee wate aate ork eee eke 14 RirGrad IR eIp IS SIR NGL eer es Sen es a A ae al Pa ite to ele hin ene deans eed ey emer erect 15 Momeceanes area andiPopulatun: Density. rx oie in 308 taggin ce yi crepe mate eye sh vor aunty 15 eo Oe ene ae er aay aey ah t CRMADAT INNIS EL BUTT Fad my BARNS Bi Poe TTT) Shas ee te cca Oe ciel Bt dome made ee ae 15 BTedatonsete ea eee ee Nh rami ne oF. Wn! Soba ae ct, Uh! ee, ae Rd eee Wp tte sce Saami 7 ii EeraaOMM CRA a oer Bort, ea at hod bo Aline digltn a at aMlalane se lie BRT Cele me oe eh sean pais * poeta: 18 REPROD UCHI ON eRe nae eee eeeiet es a er alanee es DNe tme te tie acous se canneekeoe acl emere pee ee mad crete! emeiea ers 18 CIEE SOT LIEICe Been cps chou WARS thie maitre Na tlh otha Sit er Bett Med AO ede i ci ae rk ct Rene Moe Se amr i iM 18 SET IMs Ba ITA Ci OG tee SO Se NP Donn nce ee ae Sere Somer ere a eran ens Cranes 19 MemppagMctive TAMmALOMiy NOLES: S05 5 cca's saline ow Sie Mid be, oa wal ened we oeyple Go eile ley eae om tae eee nore 19 Berrie e EIEN yale meen itn ea =e ih ela, Selene ced We std) ee ems lan pr ot pO aay © eens alee aeieene ertoc sseeearer 20 ROCHON CIN PICOM err nae Sen ener ce na dins. Sores Gus igh ha ok Orestes, a NSW iaae Na aye Keeg raile € sonata erent Chane 20 SMM NDR eh Pe Teng he ede ce ri te iy aoe Ate Coie e eno ratnr = Selst aah g he ate ela 21 Male Doemimnance Rank.and Reproductive Success.) 5" 17 fe PO ee es es 2A RapeceainpAOULDLECA Rei sete ae See Ler Me eR A RS PRUE PRR ood nae een ieee 21 SECT a hss | RR Se Se Bae de SD ae es SR ar ge A a Si tt 22 PP ATTUUTI GLO TIS ee ee a ERE Hoel Nees eta AE Meee cet ala Onane GAG Way eriake ene or ete ALOE Reta su Taps PARR ate eR al), FOE Oe ee a Pe Be PEE Pe Ss ee ae ee 23 PSU SE me UL a cae et ee ote Le, eae LUI BRU Ee re OE NS iat Secret tne 23 Rig anntiN eee Meee eg ge) nad, Skane GPRS oe GPa rane Rhee er onaa oh ee, inate Se arene Mao 23 UA irelins here: WEOT ALby cit... Sena PW on Gt, ote Cenihs Daa ee eek. Mee Mee a em Senne a 24 WESUC Ve tees ARE BOR) De AO at Baar ae aac Ea aie hk SR ater ON ee eM Shik od Boa 25 Miglieointatlatetactons .¥. 122i. YE AE Pe Rs Oe ert UE aie ore eee 23 bangevity; Reproductive Senescence; Death Rate...) in eee ce ae i ee ee ee ee 26 Poouldtion Growin Rate PRE er Se Pe OR AE BE PEN nD sla he we ate one ove 26 eee rae ae MA ALES TAY, Se ee eR a REO OS RECUR ROLE ROTEL NE Se pth Malate ae Soke Stee cee be 27 SPEED Le ile aene Se es A oie ee EE) Be Oa a ROL OP Og OOM CO rs cM A ke a i as ee 28 UCTS 08 (eal Mau gS ae eh eA SE A Re a ie ae ae See ene ie eae be 28 ire WSenies es es iene ae OR a rcek. Sone Ne wie ere ein y Sete Min) his ers ty ee he 30 Apmis SMUT tek Regs Se Ne SENS ellen ah ota ee Hehc kate Ta nata hated a ee Me ee tote is Mea nce mrs eee 30 DISET APUG TN aks eee eae ee ee te AR Aas Ute EATEN LEE AD iy cea ahaa s Moma Serene aa 30 NFO GIS om, << to Uatek se apm Ee Mee A ee Set RN SSeS Moe eee Rae Te GON kite ane ty ar mnaten sl 30 Rear s oe. 35.50 A OO MAI es PR eae). ee eee Pee 30 Specimens xaminedinn 2 ety th wep eo amie Mate acer Baten a cee eel = 31 EV ORUMONVANDsIDISPERSATE Aa ae: Fre Ete aan eee rete see il rte SPE Pat Ale ledaar t Eo Gee Rava, oct oleae et che Mane ete etts\ = Ji PERN EDG MEN TRUSTe trae a ed Oe ele eT) Se Re a Pe Oe hee 32 Le GARAGE © uD eee eee ee oe Au 0s, ics anc eile Rdeerene c Mae ec erect ee cut ane ine! sueseaauths 32 Aprenpix I: SpecIMENS EXAMINED (DETAILS). 2... «oo 6 ss ele nce ces ee opine » nao» he eniee eat 42 APppENDIx II: GAZETTEER OF MACACA SYLVANUS LOCALITIES «os 0006.0 5.5 sds 6 58 ay oles 48 6 6 es ge 42 AppENDIX III: CENTRUM LENGTH AND VERTEBRAL COUNT OF CAUDAL VERTEBRAE IN MACACA SYLVANUS . . .52 AppeNDIX IV: Group SIZE AND ADULT SEX COMPOSITION IN MACACA SYLVANUS POPULATIONS. .......+55 a3 AppENDIX V: COORDINATES, GEOLOGICAL AGE, AND REPORTED IDENTIFICATION OF CIRCUM-MEDITERRANEAN MACADUR FOSSIES, 5. ocd wien Wood a 0% % ie Owe cepa hee ela eal nein pea 54 List of Illustrations 1. Map of northwestern Africa showing extant and Holocene localities of Macaca sylvanus ......... 3 2. Adult pelage in Macaca sylvanus A Wale: 60-10 Ol 55. oa snes os eon elle ee ne pw he Buh WO) Lk 4 @ 8 808 0 290 dye Sale de pen 5 a ae OC i a i ere aR eC 5 C Hstrous female. post-molt™...b.s ec ss Mec ele wee ie ee ok Se ae ea pee be tn oer fe 3. Tail vestige in Macaca sylvanus . 0.0.00 ee te ne ete wae ne wth eel yk Sr 7 4. Ventral view of sacrum and caudal vertebrae in Macaca sylvanus .. 1.0... cc eee eee ees 9 5. Skull of adult female Macaca sylvanus ... 25. ccc ce ee ee eet we * > Gee 10 6; Skull of adult male Macaca sylvanus:.. . «2. 00... 5 v0 2 + «jeloenld «yee nh anes a wes peg eee 11 7. Phylogenetic relationships among Moroccan and Algerian populations of Macaca sylvanus as indica- fed by MIDNA haplotypes 2 cos os). x cue tise sas nt so ee oo net pe oes el rn 13 8. Copulation posture in Macaca sylvanus ... 1.6 oc te ee ees one os bie 22 9. Geographic distribution of circum-Mediterranean macaque fossils compared with geographic distribu- tion of extant populations and Holocene extinct populations of Macaca sylvanus ...........+- 28 List of Tables 1. Frequency distribution of elevation records of Macaca sylvanus ... 1... 660 eee 4 2. External measurements and proportions in wild-collected Macaca sylvanus .... 2.2.66 00 40+ e eee 6 3. Tail length and relative tail length in Macaca sylvanus specimens collected near Azrou, Moyen Putian WRT E ol, 22 3 os Bees ek tah P Sse J ea: aoa ge oe bea dee Beem ca Depa be ot wr co oe 8 4. Cranial measurements and proportions in wild-collected Macaca sylvanus .... 1.2.66 0040 eevee 12 5. Mitochondrial DNA haplotype frequencies in Gibraltar and North African samples of Macaca sylvanus. . 12 6. Estimated dates of divergence of Macaca sylvanus sister groupS .... 2... 6-0 eee eee ee ees 13 9° Habitats of Macaca Sylvanus 25 5.3. [eel E Doe on 6 es 6 6 ohh a Side fa when, MO ee 14 8. Frequency of terrestriality in Macaca sylvanus populations. .......... 0060 e ee eee eee eee 15 9. Home range area in natural populations of Macaca sylvanus .... 2... eee es 16 10, Population density in Macaca sylvanus.. .. . 06... 6 isk cid s Henin os bo cae 16 11. Number of gymnosperm and angiosperm species consumed by Macaca sylvanus populations. .... . 17 12. Allocation of feeding time to various dietary components by Macaca sylvanus groups in Grande gf ee oy ee Lue See ORIOL Ir ry SCN 18 13. Seasonal mating periods and birth periods reported in Macaca sylvanus populations............ 19 14. Mean annual birth rate in Macaca sylvanus groups . .. 2 6 000 ee eile ee es Sl 23 15; Neonatal sex ratio in: Macaca sylearius: i552. 5 isos ejn' ess on oye so aa Ge ee os wie rr 24 16. Twinning frequency reported in semi-natural populations of Macaca sylvanus ..........+.++5: 24 17. Stillbirth frequency in semi-natural populations of Macaca sylvanus .. 1... 060 ee a 18. Infant mortality rate in natural and semi-natural populations of Macaca sylvanus ..........++. 2s 19. Mean annual death rate in semi-natural populations of Macaca sylvanus... 1.6.0 060 es 27 20. Demographic growth rate in natural and semi-natural populations of Macaca sylvanus.........-.- 27 21. Evolution and dispersal of circum-Mediterranean macaques: hypothetical chronology........... | Systematic Review of the Barbary Macaque, Macaca sylvanus (Linnaeus, 1758) Jack Fooden Abstract The Barbary macaque, Macaca sylvanus (Linnaeus, 1758), is systematically reviewed, based on a study of 103 specimens and survey of relevant literature. This review includes analyses of external characters, cranial characters, DNA variation, and karyology. Information also 1s presented concerning natural history, reproduction, fossils, and taxonomic history. Current knowledge of geographic variation in M. sy/vanus is insufficient to warrant recognition of subspecies. Although M. sy/vanus is now restricted to northwestern Africa, fossil evidence suggests that this species is a relict of the ancestral macaque stock that originated in Africa ca. 7-6 million years ago (Ma) and dispersed to Eurasia ca. 6-5 Ma. Macaques formerly were widely distributed in Europe, but became extinct there ca. 0.100—0.020 Ma; macaques have survived in Asia and are now represented there by ca. 20 species. An annotated gazetteer lists 166 localities where M. sy/vanus has been collected, observed, or reported. Introduction The Barbary macaque, Macaca sylvanus, 1s unique within its genus in two obvious respects: (1) it is the only macaque species that is distributed outside of Asia, and (2) it is the only macaque species in which the tail is reduced to an inconspicuous boneless vestige or is completely absent. Probably as a result of its juxta- Mediterranean distribution and its distinctive tail reduction, the Barbary macaque was well known to ancient Phoenicians, Etruscans, Egyp- tians, and Greeks—including Aristotle (Mc- Dermott, 1938, pp. 28, 29, 38, 88; Goudsmit & Brandon-Jones, 1999, p. 49). This species is the first primate listed in Gesner’s (1551, p. 957) classic Historiae Animalium, from which it was cited by Linnaeus (1758, p. 25). Division of Mammals, Department of Zoology, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605-2496, U.S.A. Materials and Methods This review of M. sy/vanus is based on study of 103 specimens (Appendix I) and review of relevant literature. ArcView GIS was used to prepare maps, and Microsoft Excel was used to calculate summary statistics. Reported cranial measurements (Table 4) are defined as follows (cf. Fooden, 1969, p. 40): Greatest length of skull—Most anterior point on rostrum to most posterior point on skull. Postrostral length— Most inferior point on either orbital margin to most posterior point on skull. Rostral length— Most anterior point on rostrum to most inferior point on either orbital margin. Zygomatic breadth—Distance between most lateral points on zygomatic arches. Names of institutions in which cited specimens are preserved are abbreviated as indicated below; the number of M. sy/vanus specimens studied at each institution is indicated by a parenthetical notation: American Museum of Natural His- tory, New York (7) AMNH FIELDIANA: ZOOLOGY, N.S., NO. 113, OCTOBER 26, 2007, PP. 1-58 l BM(NH) The Natural History Museum, London (8) CMNH Carnegie Museum of Natural His- tory, Pittsburgh (1) FMNH Field Museum of Natural History, Chicago (4) IRSN Institut Royal des Sciences Natur- elles de Belgique, Brussels (17) ISR Institut Scientifique, Rabat (7) MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (2) MNHN Muséum National d’Histoire Nat- urelle, Paris (10) NHMBa Naturhistorisches Museum, Basel (1) NHMBe Naturhistorisches Museum, Bern (1) NMS Natur-Museum Senckenberg, Frankfurt (4) RMNH Nationaal Naturhistorisch Muse- um, Leiden (10) USNM National Museum of Natural His- tory, Washington, D.C. (19) ZMB Museum fiir Naturkunde, Berlin, (3) ZMUZ Zoologisches Museum der Univer- sitat Ziirich (4) ZSBS Zoologisches Staatsammlung, Mu- nich (5) Geographic Distribution Extant natural populations of M. sylvanus are disjunctly distributed in the Atlas and Rif Mountain ranges in Morocco and Algeria (Fig. 1); latitudinally the geographic range ex- tends from ca. 31°15'N to 36°45'N, and longitu- dinally it extends from ca. 7°45’'W to 5°35’E. Moroccan and Algerian populations are separat- ed by a gap that is ca. 700 km wide, and narrower gaps separate disjunct populations within these two countries. Holocene extinction records (Fig. 1, 28 populations) indicate that distribu- tional gaps were narrower within historic times and further indicate that the range of the species recently extended east of Algeria to ca. 10°E in neighboring Tunisia; an early reference to M. sylvanus in Tunisia has been provided by Her- odotus (ca. 484-425 B.C.E.) (McDermott, 1938, p. 56). The known elevational distribution of M. sylvanus is 400-2300 m (Table 1); half of the 62 available elevational records are at or above 1800 m. No other nonhuman primate is known to have ever been sympatric with M. sy/vanus. The population of M. sylvanus that currently inhabits Gibraltar apparently is not native. There is no mention of this species in ancient commentaries on Gibraltar (Joleaud, 193la, p. 138; McDermott, 1938, p. 56). Instead, the oldest known reference to monkeys on Gibraltar is in an 18th century document preserved in the British Museum (Sayre, 1862, p. 454; cf. Buffon, 1789, p. 36); this document indicates that “a great quantity” of Barbary macaques were introduced into the British garrison in 1740 (cf. Busk, 1877, p. 129; Trouessart, 1905, p. 359; Morris & Morris, 1966, p. 15; Fa, 1981, p. 74). Additional introductions occurred in 1813 and 1860 (Fa & Lind, 1996, p. 235), but the population in 1925 was reduced to three indi- viduals (Kenyon, 1938, p. 115), and in 1938- 1945 major restocking occurred (Fa, 1984b, p. 266). A recent study of mtDNA variation in the existing Gibraltar population indicated that this population includes individuals of both Moroccan and Algerian ancestry (Modolo et al., 2005, p. 7397). Lydekker (1893-1894, p. 118) and Forbes (1894, p.5) indicated that during the 19th century, M. sylvanus inhabited southern Spain in the vicinity of Gibraltar. This appears to have been an error (cf. Kelaart in Newman, 1846, p. 1292; Joleaud, 193la, p. 138), and, in a sub- sequent publication, Lydekker (1916, p. 7) omit- ted Spain from the species distribution. An introduced population of M. sy/vanus was recently established in Morocco in 1985, when more than 40 formerly captive individuals were deliberately released near Nador, a northern coastal town at ca. 35°00'N, 3°00'W (Aulagnier & Thevenot, 1986, p. 56; cf. Chivers, 1984, p. vii; Anonymous, 1988, p. 50). Total Population Estimates In 1984, Fa et al. (1984, pp. 86, 91, 99, 101) estimated the total extant natural population of M. sylvanus to be 14,000—23,000 individuals (9,000-17,000 in Morocco, 5,000—6,000 in AlI- geria). In 1992, Lilly and Mehlman (1993, p. 327; cf. Mehlman & Lilly, 1993, p. 334) indicated that the population had declined to 10,000—16,000, and, in 2005, Modolo et al. (2005, p. 7392; cf. FIELDIANA: ZOOLOGY 5 ro } } 7 ~ SPAIN 60 53 See aaB3 O 62 * = 56 gS 61 OQ 468 4, 49 bd 55 = 45 47 1 ° 59 We ALGERIA | TUNISIA e Extant populations © Holocene extinctions e Introduced populations 0 100 200 300 Kilomete —k— 0° 5° {= Fic. 1. Map of northwestern Africa, showing extant and Holocene localities of Macaca sylvanus; for documentation, see Gazetteer, Appendix II. Key to Locality Numbers: MOROCCO and U.K. 1. Goundafa. 2. Ourika, Jebel; Ourika, Oued: Tourchte; Zaclia Si Fatma; Zaclia Si Fatma, S of. 3. Ouzoud, Cascades d’. 4. Ait Mehammed, S of. 5. Tillouguit N’Ait Isha, S of. 6. Ouaouizahrt, MF de. 7. Ighrem, Jebel. 8. Asker, Zaouia; Idrous, Jebel: Kousser, Massif du; Tsemair. 9. Ait Sokhmane forest; Azrar; Boutferda vicinity; Haounet, Forét d’El; Ijbertene. 10. El Kisba vicinity; Komuch. 11. Asserdoun. 12. Khenifra area. 13. Ajdir, Plateau d’; Ali ou Zrou, Jebel; Annocer; Arhbalou N’Irkraouen; Azigza, Aguelmane; Bou Ceédre; Bou Igaouerh, Jebel; Imi Ourarn; Kerrouchene; Khenifra, 22 km E; Mimejad; Senoual region; Tafechna; Tiguerouguine. 14. Abid, Oued el, gorge; Arhbala, N of; Tizi M’Isly, N of. 15. SidiYahya ou Youssef, 8-10 km W of; SidiYahya ou Youssef, S and W of. 16. Tounfit, 15-20 km SSE of. 17. Tounfit, N and W of. 18. Tagoulelt vicinity. 19. Itzer, 10-15 km E of; Itzer, W of. 20. Assaka N’Ouam; Zad region. 21. Seheb. 22. Afennourir; Ain Kahla; Ain Leuh; Azrou, 18 km S; El] Hammam; Ouiouane; Oum Er Rbia, Oued, source; Sidi Mguid vicinity; Zaouia de Ifrane. 23. Azrou; Azrou, 5 km $; Azrou, above; Azrou Forest; Azrou region; Bou Jirirh; Cédre Gouraud; Ifrane; Michlifene; Ras El Ma. 24. Imouzzer du Kandar, vicinity. 25. Taffert; Tahar Souk, N of. 26. Bou Iblane, Jebel; Tamjilt, below. 27. Tahafourt. 28. Rhiata mountains. 29. U.K.: Gibraltar; Gibraltar, N end of mountain. 30. Moussa, Jebel. 31. Fath Lemhar. 32. Ahfa- Timesh; Buzeitune, Jebel; Derti-Na, N of; El Haouz region; Kelti, Jebel; Sidichmim; Tétouan vicinity; Zarka cascade vicinity. 33. Bouhassim, Jebel. 34. Adelma, Oued; Beni Mohammed; Chechaouene, Jebel; El Kelaa; Farda, Oued; Kaiat, Jebel; Lakraa, Jebel; Magoo, Jebel; Sidi Salah, Jebel; Sidi Siah; Talaat, Ardhousse; Talassemtane; Tasaot, Jebel; Tilljida, Oued; Tisouka, Jebel. 35. Tisirene, Jebel. 36. Targa vicinity. 37. Besene, Bab. 38. Ketama vicinity. 39. Tidirhine, Jebel. 40. Tizi Ifri. 41. Al Hoceima. 42. Beni Touzine vicinity. 43. Nador vicinity. ALGERIA. 44. Rond-Point des Cédres. 45. Tighret forestry post. 46. Chiffa, Gorges de la. 47. Chiffa, Gorges de la, ca. 15 km S of Blida. 48. Chrea. 49. Bou Zegza, Djebel. 50. Palestro, Gorges de; Tala Kitane. 51. Ait Oubane, Forét d’; Cheminot; Djurdjura, Djebel; Guessig, Mount; Hotel; Icetcifene; Ras Timedouine; Source des Singes; Tala Gulief; Tala Rana; Tigounatine; Tikjda. 52. Akfadou, Forét de; Beni Rhobri, Forét des; Taourirt Irhil, Foret de. 53. Bouak, Cap-Cap Carbon, between; Pic des Singes National Park. 54. Akra, Chabet El; Borj Mira; Darguinah; Kherrata, Gorges de; Takoucht, Djebel. 55. Babor, Djebel. 56. Adendoun, Djebel; Dar El Oued; Guerrotich, Forét Domaniale de; Ziama Mansouria, E of. 57. Kebir, Oued El, gorge. 58. Constantine. 59. Collo, Kabylie de. 60. Stora, “Oran”; Stora vicinity. 61. Azzaba, W of. 62. Edough, Djebel. TUNISIA. 63. Tabarqah. 64. Zaghwan. Camperio Ciani & Palentini, 2003, p. 63) re- ported a further decline tol0,000. The main factor responsible for population decline appar- ently is anthropogenic destruction of habitat (Fa, 1986a, p. 250; Camperio Ciani et al., 2005, p. 640; Waters et al., 2007, p. 107). M. sylvanus has been designated as Vulnerable on the IUCN Red List of Threatened Species (IUCN, 2006). Pelage (Fig. 2) In M. sylvanus adults and subadults, prime pelage coloration on the dorsal surface of the trunk typically is strongly variegated (Fig. 2C), pale (pale buffy to golden brown to burnt orange) and dark (blackish). This variegation is a result of conspicuous banding of individual dorsal hairs: in one adult male (UsNM 255979, FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 3 Tas_e |. Frequency distribution of elevation records of Macaca sylvanus (see Gazetteer, Appendix I1).! Elevation (m) Number of records 400-599 600-799 800-999 1000-1199 1200-1399 1400—1599 1600-1799 1800-1999 2000-2199 2200-2300 Total to NK ORF DAHhO~AN—NN on ' Where elevations were originally reported as a range of values, the midpoint of these values is tabulated here. Gibraltar, zoo specimen), interscapular hair length is ca. 80 mm, the basal 50 percent of each hair is dark brown, the middle 30 percent is pale buffy, and the distal 20 percent is blackish; in other adults and subadults, the pale intermediate band often is more brightly colored. Premolt pelage is faded and drab (Fig. 2A). The pelage in one exceptional adult or sub- adult female (USNM 258206, Gibraltar, zoo specimen) is strikingly erythristic—reddish brown on crown, trunk, and appendages. Mor- occan specimens that apparently are similarly erythristic have been reported by Panouse (1957, p. 25: “franchement au roux”) and Aulagnier and Thevenot (1986, p. 56: “brun-roux sur le dos”). According to Bédé (1927, p. 28), speci- mens obtained in Morocco were generally more erythristic than those obtained in Algeria, but the purported geographic difference remains to be adequately documented (Didier & Rode, 1936, p. 13; Rode, 1937, p. 158). Pelage on the crown is variegated like that on the back, but crown hairs tend to be more brightly colored. The sides of the face are bounded by streaks of dark brown hair, whereas the face itself is thinly haired and lightly pigmented. The lateral facial crests in 13 speci- mens examined are restricted to the posterior mandibular region (infrazygomatic crests; Foo- den, 1995, p. 19). On outer surfaces of the pectoral and pelvic appendages, variegation tends to decrease from proximal to distal segments, generally becoming pale brown to medium brown on dorsal surfaces of the hands and feet. The ventral surfaces of the trunk and appendages are thinly haired, pale buffy to whitish. At birth, dorsal pelage is blackish, and ventral pelage, which is thin, is slightly paler (Fa, 1984c, p. 338; cf. Carpentier, 1931, p. 275; Deag & Crook, 1971, p. 188; MacRoberts & MacRo- berts, 1971, p. 39). The gradual transition from blackish infantile pelage to variegated brownish juvenile pelage begins about age 45 days and is completed by about age 145 days. Pelage length in juveniles is less than in subadults and adults, but the coloration is generally similar. Molting occurs during spring and summer and usually is complete by September or October (Fig. 2C; cf. MacRoberts & MacRoberts, 1971, p. 39; Merz, 1976, p. 59; Deag, 1984, p. 124). In females with new infants, molting usually begins about two months after parturition (Fa, 1984¢c, p. 339). The bright new fur appears first on the head, arms and forearms; subsequently, it appears more caudally and ventrally. External Measurements and Proportions Head and body length measurements are available for a small sample of wild-collected M. sylvanus specimens, all of which originated near Azrou, Moyen Atlas, Morocco (Table 2). In this sample, mean head and body length is 556.8 mm in four adult females and 634.3 mm in three adult males; the female/male dimorphism ratio is 87.8%. Sexual dimorphism of head and body length is similar in six captive specimens or specimens of unknown history—573.0 mm in four adult females (AMNH_ 19014, BM[NH] 11.11.11.1, usnm 258206, 502460) and 630.5 mm in two adult males (Cabrera, 1932, p. 201; mMcz 15296); in these specimens, the female/male length ratio is 90.9%. No information is available concerning geographic variation of head and body length in M. sylvanus. Mean body weight in wild-collected adults is 9.9 kg in 18 females and 14.5 kg in 33 males (Table 2; cf. Roberts, 1978, p. 233; Fa, 1984c, pp. 341-346; 1989, p. 58); the female/male di- morphism ratio for this volumetric measurement is 68.3%. Mean weight in four adult females collected in Morocco (9.2 + 0.82 kg) is not significantly different (P > 0.05, t-test) from that in 14 females probably trapped in Algeria (10.1 + 1.03 kg); weights of Moroccan adult males are unavailable (cf. Table 2, footnote 4). FIELDIANA: ZOOLOGY Fic. 2. Adult pelage in Macaca sylvanus. A. Male, premolt. B. Male, molting (note new pelage on head and shoulders). C. Estrous female, postmolt. Photographed 29 September 2006 at Cédre Gouraud, Morocco, by G. Fooden; image in figure B is photographically reversed. Relative to adult size in M. sylvanus, ears in foot in infants is only ca. 60% of that in infants are larger than hind feet (Table 2). adults. Length of the ear in infants is ca. 80% of An inconspicuous vestigial tail is variably that in adults, whereas length of the hind present in M. sylvanus (Fig. 3; Table 3; cf. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 5 TABLE 2. . . . ° External measurements and proportions in wild-collected Macaca sylvanus.'* Relative hind Relative Head and body Weight Hind foot foot length Ear length ear length Age/sex class* length (mm) (kg) length (mm) (HF/HB x 100) (mm) (E/HB x 100) Infants 266.5 1.0 101.5 38.2 40.0 15.0 246-287 0.9-1.0 96-107 37.3-39.0 36-44 14.6-15.3 (2) 2 (2) (2) (2) (2) Juveniles 521.0: = 28560 “eS 159.0 = 13:89 30.5 + 1.10 45.0 + 6.24 8.6 + 1.00 500-553 6.2-8.6 150-175 29.4-31.6 38-50 7.5-9.4 (3) (3) (3) (3) (3) (3) Subadult males 604.0 559-649 182 28.0 51 7.9 2 : (/) (/) (J) (/) Adult females §56:8 = Lea 0 E04 168i 171 30.1 + 0.68 48.8 + 1.26 8.8 + 0.10 540-566 8.0—12.0 166-170 29.3-30.9 47-50 8.7-8.9 (4) (8) (4) (4) (4) (4) Adult males 634:3%= 25.725 4 17S 179.0. 71.94 Pi WS ageing left SS 1S 8.1 + 0.42 605-653 10.0-18.0 170-185 26.4-30.1 50-52 7.8-8.6 (3) (33) (3) (3) (3) (3) ' Mean + SD (where n > 2), extremes, and sample size (italicized figures in parentheses). > Most measurements are of specimens collected near Azrou, Morocco (BM(NH), USNM). Exceptions are weights of 14 adult females and 33 adult males that were reported by Delson et al. (2000, pp. 144-145); these 47 adults probably were trapped and weighed in Algeria (cf. Scheffrahn et al., 1993, p. 383; 1996, abstract no. 240). Measurements of USNM specimens are from specimen tags and collectors’ field catalogs. * Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, M3 in females or C in males incompletely erupted; adults, all permanent teeth completely erupted. * Excludes USNM 476781 (juvenile) and UsNM 476791 (weight recorded as ‘‘>10 kg’’) (cf. Delson et al., 2000, p. 144). Lydekker, 1893-1894, pp. 117-119); Aristotle (d’A. W. Thompson translation, 1984, p. 799) accurately described the tail vestige in this species as “*... a very small one—a sort of hint of a tail.” In a series of 13 Moroccan specimens that were carefully examined by the collectors, a tail vestige, varying in length from 4mm to 22 mm, was detected in eight individuals (62%); although the sample size is small, the data suggest that the tail vestige might be more frequent in males (86%) than in females (33%). Tail vestiges of similar length are known to occur in three captive males (Hill, 1974, p. 622, 18.7 mm; FMNH 60716, 9mm _ [Fig. 3]; Mcz 15206, 22 mm). Relative tail length probably is greater in infants than in postinfants (Table 3), which suggests that the postnatal growth rate of the tail is less than the postnatal growth rate of the head and body. Caudal Vertebrae Caudal vertebrae also are vestigial in M. sylvanus (Fig. 4), and, as previously noted by Waldeyer (1896, p. 783) and Schultz (1925, pp. 249-250), these vertebrae do not extend into the external tail; for this reason, Waldeyer and Schultz have characterized the tail vestige in M. sylvanus as “soft.” The number of caudal vertebrae in this species varies from two to five (Appendix III), with a modal number of three (13 of 29 specimens reported). Reduction in number of caudal vertebrae in M._ sylvanus apparently exceeds that in humans, where the modal number of caudal vertebrae is four (Schultz & Straus, 1945, p. 613). Cranial Characters Mean greatest length of skull in wild-collected M. sylvanus is 118.7 mm in four adult females and 135.5 mm in five adult males (Figs. 5, 6; Table 4); the female/male dimorphism ratio is 0.88. Available evidence does not permit analysis of geographic variation of skull size in this species (cf. Table 4, footnote 4). In wild-collected M. sylvanus, mean relative zygomatic breadth increases from 60.9% in FIELDIANA: ZOOLOGY 10mm Fic. 3. Tail vestige (indicated by arrow) in Macaca sylvanus (FMNH 60716, infant male, zoo specimen); tail length is 9 mm. (Photograph by R. Banasiak.) infants to 69.2% in adult males, and mean rostral_postrostral ratio increases from 23.8% in infants to 59.0% in adult males (Table 4). As in other macaques, postnatal relative growth of the rostrum greatly exceeds that of the zygomatic arches. A sagittal crest is relatively rare in M. sylvanus. In adult male skulls examined, this crest is absent in four wild-collected specimens and present in only two (mcz 15296, MNHN CG 1831/835) of nine captive specimens or specimens of unknown history (overall frequency, 2/13 = 15.4%). FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE t Taste 3. Tail length and relative tail length in Macaca sylvanus specimens collected near Azrou, Moyen Atlas, Morocco; measurements from specimen tags and collectors’ field catalogs. i Head and body Tail length” Relative tail length USNM No. Age! Sex length (mm) (mm) (T/HB < 100) 476788 Infant Male 246 15 6.1 476789 Infant Male 287 15 EZ 476790 Juvenile Female 510 0 Be 478329 Juvenile Female 500 15 3.0 476781 Juvenile Male 553 15 Ze 476784 Subadult Male 649 pHs 3.4 476782 Adult Female 561 0 — 476783 Adult Female 566 4 0.7 476786 Adult Female 540 0 —— 476787 Adult Female 560 0 — 476780 Adult Male 605 20 33 476785 Adult Male 653 22 3.4 476791 Adult Male 645 0 —— ' For dental specifications, see Table 2. 2 Cf. MacRoberts and MacRoberts, 1971, p. 39; Burton and de Pelham, 1979, p. 109 (apparent decimal point placement errors in this publication; tail length reported as “0.5 mm” and “0.53 mm” in two infant specimens). Based on analyses of 27 cranial dimensions, Pan and Oxnard (2004, p. 367) have reported that cranial differences between M. sylvanus (21 adult specimens) and 10 Asian macaque species (303 adult specimens) greatly exceed correspond- ing differences among the 10 Asian species. Unfortunately, the provenance of the M. sylva- nus specimens (and non—M. sylvanus specimens) studied in this research project is not specified (cf. Table 4); some artificially provisioned indi- viduals might have been included (cf. Fooden & Aimi, 2005, p. 17). Molecular Biology and Genetics DNA Variation Virtually all studies of interspecific DNA variation within the genus Macaca have in- dicated that M. sy/vanus is the sister group of Asian macaque species (Hayasaka et al., 1996, p. 1050; Morales & Melnick, 1998, pp. 15-16; Deinard & Smith, 2001, p. 50; Tosi et al., 2003, pp. 1427, 1430; Modolo et al., 2005, p. 7394; Liu & Pearl, 2007, p.411; Ziegler et al., 2007, p. 811). A recent exception is Tosi et al.’s (2003, p. 1426) study of Y-chromosomal DNA, in which the phylogenetic relationship of M. sylvanus was unresolved. The most comprehensive survey of intraspe- cific DNA variation in M. sylvanus has been published by Modolo et al. (2005, p. 7393). In this study, a 468-bp sequence of hypervariable region | of the mtDNA control region was analyzed in 68 samples of Gibraltar monkeys (artificially introduced population; see above, Geographic Distribution), 116 samples of Mor- occan monkeys (3 localities), and 96 samples of Algerian monkeys (4 localities) (Table 5). Of the 24 haplotypes detected in non-Gibraltar samples, nos. 1-15 were restricted to Morocco and nos. 16-24 were virtually restricted to Algeria (cf. Pastorini et al., 2000, p. 166; Crouau-Roy et al., 2001, p. 157; von Segesser, 2002, p. 161); the sole exception is haplotype no. 16, which was detected in 25 Akfadou (Algeria) samples and four Rif (Morocco) samples. Modolo et al. (2005, pp. 7394, 7397) plausibly concluded that the anomalous geographic distribution of hap- lotype no. 16 probably is a result of Algerian monkeys having been artificially transported and released in the Rif mountains. Haplotype evi- dence (Table 5) also indicated that the Gibraltar population has resulted from artificial introduc- tion of monkeys from the Rif (Morocco) and Akfadou (Algeria) regions (Modolo et al., 2005, p. 7397): Phylogenetic analysis indicates that Algerian mtDNA haplotypes are more divergent than Moroccan haplotypes (Fig. 7); in fact, Algerian haplotypes at Pic des Singes (haplotype no. 18) and Kherrata (haplotypes nos. 19-22) are more similar to Moroccan haplotypes than they are to other Algerian haplotypes (cf. Scheffrahn et al., FIELDIANA: ZOOLOGY Fic. 4. Ventral view of sacrum and caudal vertebrae (Cd 1—Cd 3) in Macaca sylvanus (FMNH 47398, adult male, ZOO specimen); vestigial Cd 3 is asymmetrically fused to Cd 2. (Photograph by J. Weinstein.) 1996, abstract no. 240). Modolo et al. (2005, changes that resulted in periodic forest fragmen- p. 7396) have indicated that the present disjunct tations and interconnections. Based on fossil clustering of mtDNA _ haplotypes could be evidence and the date of the Messinian Salinity a product of Pleistocene and Holocene climate Crisis (Delson, 1980a, p. 16; Tosi et al., 2003, FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 9 Fic. 5. p. 1424), Modolo et al. (2005, p. 7393) have postulated that M. sylvanus originated ca. 5.5 million years ago (Ma) and have inferred that extant clades and subclades of M. sylvanus 10 Skull of adult female Macaca sylvanus: USNM 476782, Azrou, 18 km S of, Morocco. (Photographs by E. M. Santana.) differentiated within the past 1.85 million years (Table 6). Microsatellite variation within and between four isolated Algerian populations of M. sylva- FIELDIANA: ZOOLOGY Fic. 6. Skull of adult male Macaca sylvanus: USNM 476785, Azrou, 18 km S of, Morocco. (Photographs by E. M. Santana.) FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 1] Taste 4. Cranial measurements’? and proportions in wild-collected Macaca sylvanus; except for one adult male obtained at Gorges de la Chiffa, Algeria, all specimens were collected near Azrou, Morocco (see footnotes 4 and 5). Greatest length Relative zygomatic Postrostral Age/sex class* of skull (mm) breadth (ZB/GL x 100) length (mm) Infants 80.3 60.9 fb S| 77.6-83.0 60.4-61.4 68.4—73.7 (2) (2) (2) Juveniles 113.0 66.6 82.6 109.8—116.3 65.7—-67.5 81.2-83.9 2 (2) (2) Adult females 118.7 + 5.48 69.4 + 2.054 85.9 + 1.66 113.7—125.7 68.2—72.4 84.1-87.5 (4) (4) (4) Subadult males 124.3 67.6 90.7 118.2-130.5 66.0-69.2 (1) 2 (2) Adult males 1955: 207° 69.2 + 1.00° 91.7 + 0.95 132°7-137-7 68.0—70.5 90.5-92.7 (5) (5) (4) ' For definition of measurements, see above, Materials and Methods. Rostral-postrostral ratio (R/PR X 100) 23.8 23.3-24.4 (2) 48.0 43.7-52.2 (2) 49.6 = 342 47.4-54.2 (4) 56.5 (/) 59.0 + 2.47 55.6-61.6 (4) > Mean + SD (where sample size exceeds two), extremes, and sample size (italicized figures in parentheses). * Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, M3 in females or C in males incompletely erupted; adults, all permanent teeth completely erupted. * 135.3 + 2.58 mm in four adult males collected in Morocco; 136.2 mm in one adult male collected in Algeria (measurement of Algerian specimen provided by D. Brandon-Jones). > Zygomatic breadth of Algerian specimen estimated by D. Brandon-Jones. nus was studied by von Segesser et al. (1999, p. 436). In a sample of 159 monkeys, six of the seven loci tested were found to be polymorphic, with 4-12 alleles per locus. Populations and population subgroups were genetically differen- tiated with respect to these loci. Genetic distance was significantly correlated with geographic distance within a population—but not between populations. Crouau-Roy et al. (2001, p. 157), who also studied microsatellite variation in M. TABLE 5. sylvanus (Modolo et al., 2005, p. 7393). Locality N DPa2 3 45-67-89 10 Tei ts 14 1S) 56: 17 U.K. Gibraltar 68 24 8 36 Morocco Rif 28 4 20 4 Moyen Atlas 69 4 16 3 ie LO) DG: 2 oS) Haut Atlas 19 6 Cues 4 Algeria Akfadou 25 23 Djurdjura 54 Pic des Singes 12 hese Kherrata 5 12 sylvanus, found 12 alleles specific to Moroccan samples and 21 alleles specific to Algerian samples. Karyology The chromosomes of M. sylvanus have been studied—with partly divergent results—by Stan- yon et al. (1980, p. 150), Dutrillaux et al. (1982, p. 97), Tihy et al. (1996, p. 10), and Morescalchi Haplotype code numbers 18 Mitochondrial DNA haplotype frequencies in Gibraltar and North African samples of Macaca 19 20 21 22 23 24 FIELDIANA: ZOOLOGY Rif Mor. Moyen Atlas Mor. Haut Atlas Mor. Kherrata Alg. Pic des Singes (hap. no. 18) Alg. Dyjurdjura Alg. Akfadou Alg. Pic des Singes (hap. no. 17) Alg. Fic. 7. Phylogenetic relationships among Moroc- can (Mor.) and Algerian (Alg.) populations of Macaca sylvanus, as indicated by mtDNA haplotypes (maxi- mum likelihood tree, condensed from Modolo et al., 2005, p. 7394); the Pic des Singes sample includes two haplotype subgroups. et al. (1998, p. 102). Stanyon et al. (1980, p. 150) found that chromosomes in M. sy/vanus are generally similar to those in M. mulatta, except that banding patterns in four chromosomes (nos. 1, 4, 9, and 16) are polymorphic in M. sylvanus but not in M. mulatta. Dutrillaux et al. (1982, p. 107) reported that chromosomes in M. sylvanus are similar to those in other macaques (except M. fascicularis) and also are similar to those in species of Papio and Lophocebus. Tihy et al. (1996, p. 10) reported that chromosome no. 18 in M. sylvanus is homologous to chromo- some no. 13 in humans; a centromeric shift probably occurred during the course of this chromosomal transition. And Morescalchi et al. TABLE 6. relationships (Modolo, 2005, p. 7394, table 4). (1998, p. 103) reported that chromosome no. 2 in M. sylvanus has a clear band adjacent to the centromere that does not occur in other species of Macaca but does occur in species of Papio; these authors also found that human chromo- some probes nos. 7 and 21 hybridized to M. sylvanus chromosome no. 2 (cf. Wienberg et al., 1990, p. 347). As in other members of the tribe Papionini, the diploid chromosome number in M. sylvanus is 42. Natural History Habitats Habitats occupied by M. sylvanus include cedar, fir, and oak forests (evergreen oak and deciduous oak), scrub, grasslands, and rocky ridges dominated by herbaceous vegetation (Table 7); although most extant M. sylvanus populations inhabit cedar forests, this might reflect current habitat availability rather than habitat preference (Fa, 1984e, pp. 281, 284). Climate in the range of this species is Mediter- ranean, with warm dry summers and relatively wet winters (Trewartha, 1978, p. 8); at elevations above 1500 m—1.e., well within the elevational range of M. sylvanus (Table 1)—snow falls in winter, and snow accumulations can persist on mountain summits until mid-May (Fa, 1982, p. 50; Mehlman, 1984, p. 172). Alvarez and Hiraldo (1975, p. 256) have reported that mon- key populations in the Rif mountains migrate to lower elevations during winter. Estimated dates of divergence of Macaca sylvanus sister groups, based on mtDNA haplotype Sister groups (see Fig. 7) Morocco/Kherrata/Pic des Singes (hapl. Djurdjura/Akfadou/Pic des Singes (hapl. no. 17) clade (Algeria) Kherrata/Pic des Singes (hapl. no. 18) no. 18) clade (Morocco + Algeria) Morocco clade Estimated divergence dates (Ma)! 1.24-1.85 1.22—1.62 clade (Algeria) Djurdjura clade (Algeria) Akfadou/Pic des Singes (hapl. no. 17) 0.78-1.01 clade (Algeria) Kherrata subclades (Algeria) Morocco subclades Akfadou subclades (Algeria) Djurdjura subclades (Algeria) ' Million years ago. 0.064—0.359 0.110-0.282 0.070—0.078 0.070—0.078 FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 13 TABLE 7. Geographic area Morocco Haut Atlas Scrub Moyen Atlas Atlantic cedar Holm oak- Mixed cedar—oak Grassland Rif Spanish fir Atlantic cedar Holm oak~ Mixed oak Scrub Algeria “ Chiffa Holm oak~ Mixed oak ¥ Tigounatine Mixed cedar—oak~ Icetcifére Herbaceous Akfadou Mixed oak* Kerrata Holm oak? Mixed oak Dominant vegetation Habitats of Macaca sylvanus. Scientific name References! — Se 5 Cedrus atlantica 1 3o@ Quercus ilex Abies pinsapo 2576; Jee Cedrus atlantica Quercus ilex Quercus ilex 6 a 9, 10 5 9, 10 — 10, 11 Quercus ilex 6 ' Key to reference numbers: /. Deag and Crook, 1971, p. 186. 2. Taub, 1977, p. 110. 3. Ménard, 2002, p. 95. 4. Camperio Ciani et al., 2005, p. 639. 5. Alvarez and Hiraldo, 1975, pp. 254-257. 6. Fa et al., 1984, pp. 86, 100, 105. 7. MehIman, 1984, p. 165. 8. Fa, 1986a, p. 222. 9. Ménard and Vallet, 1986, pp. 173, 174. /0. Ménard, 2002, p. 95. 1]. Ménard, 1985, p. 452; 2002, p. 95. > Evergreen. * Deciduous. Terrestriality/Arboreality Macaca sylvanus spends most of the daylight hours on the ground (Table 8); reported mean frequency of daytime terrestriality varied from 68% to 83% in Morocco and from 58% to nearly 100% in Algeria. Infants and juveniles tended to be less terrestrial than adults (Merz, 1976, p. 60; Deag, 1985, p. 408; Ménard, 1985, p. 466; Ménard, & Vallet, 1986, p. 176; Machairas et al., 2003, p. 189); in populations studied at Afennourir and Cédre Gouraud (=Gouroud), Morocco, juveniles were 70% terrestrial and adult males were 81% terrestrial (Machairas et al., 2003, p. 189). Frequency of terrestriality also varied seasonally (Deag, 1985, pp. 408-409; Ménard & Vallet, 1986, p. 190; 1997, p. 292). This species flees into trees to escape danger (Merz, 1976, p. 60; Deag, 1985, p. 408), and it sleeps in trees (Taub, 1977, p. 128; de Turkheim & Merz, 1984, p. 247; Mehlman, 1989, p. 285; Hammerschmidt et al., 1994, p. 279) or in caves on rocky cliffs (MacRoberts & MacRoberts, 1971, p. 45; Alvarez & Hiraldo, 1975, p. 258; Fa et al., 1984, p. 94; MehIlman, 1984, pp. 190— 191). 14 Group Size and Composition Reported size in 68 natural groups of M. sylvanus averaged 27.1 individuals and varied from seven to 88 individuals (Appendix IV). Limited available evidence has suggested that group size might average smallest (18.3 individ- uals) in the Moroccan Rif, largest (50.2 individ- uals) in the Algerian Grande Kabylie, and intermediate (28.6 individuals) in the Moroccan Moyen Atlas (Appendix IV). Group size gener- ally is fairly stable in this species, but it was unstable in an Algerian population that inhabits a rocky habitat in the Djurdjura mountains (Ménard et al., 1985, p. 72; 1990, p. 169; Ménard, 2002, pp. 96, 98); in this marginal habitat, groups frequently split into subgroups that subsequently reunite in various combina- tions. Solitary males, as far as 7 km distant from the nearest group, have been reported in the Moroccan Rif (Mehlman, 1985, p. 204; 1986, p. 75). Provisioned semi-free-ranging groups, which sometimes have included as many as 200 members (de Turckheim & Merz, 1984, p. 243; Kuester & Paul, 1992, p. 194), apparently can become larger than natural groups. FIELDIANA: ZOOLOGY TABLE 8. Frequency of terrestriality in Macaca sylvanus populations. Mean frequency Locality Sample size of terrestriality References’ Morocco: Moyen Atlas Ain Kahla (cedar) 11,202 observations 83% 1 Afennourir (cedar) 317 hr 68% 2 Cédre Gouraud~ (cedar—oak) 267 hr 771% 2 Algeria: Grand Kabylie Tigounatine (cedar—oak) 2,587 scans 68% 3 Akfadou (oak) 2,608 scans 58% 3 Icetcifére (mountain ridge) _- ~100% 4 France: semi-free-ranging Kintzheim (mixed European forest) _- >50% 5 ' Key to reference numbers: /. Deag, 1985, p. 409. 2. Machairas et al., 2003, p. 189. 3. Ménard and Vallet, 1997, p. 292; cf. Ménard, 1985, p. 466; Ménard and Vallet, 1986, p. 176. 4. Ménard and Vallet, 1986, p. 190. 5. Merz, 1976, p. 60. ? Alternate locality name: Gouroud. Within natural groups of M. sy/vanus, the number of adult males apparently averages only slightly less than the number of adult females (Appendix IV). The known ratio of adult males to adult females averaged 0.70 in the Moroccan Moyen Atlas, 0.99 in the Moroccan Rif, and 0.84 in the Algerian Grande Kabylie. Group Fission A M. sylvanus group in the Algerian Grande Kabylie that increased in size from 38 members in 1983 to 88 members in 1988 permanently subdivided inl989 into three daughter groups (50, 24, and 13 members; | member disappeared) (Ménard & Vallet, 1993a, p. 483; 1993b, p. 105; Lathuilliére et al., 2004, p. 176). As noted above, 88 is the maximum group size known in natural groups of M. sylvanus, which suggests that fission could be inevitable in groups that approach this size. Fission also has been reported in semi-free- ranging European groups of M. sy/vanus (Prud’- Homme, 1991, p. 10; Fa & Lind, 1996, p. 237; Kuester & Paul, 1997, p. 946; Fischer, 1998, p. 800; Kiimmerli & Martin, 2005, p. 1232). Female matrilines generally remained intact during the course of these fission events, which usually began during the mating seasons. Home Range Area and Population Density Available evidence suggests that home range area in natural groups of M. sylvanus averages smallest (18.4 ha) in the Moroccan Moyen Atlas, largest (804.5 ha) in the Moroccan Rif, and intermediate (279.7 ha) in the Algerian Grand Kabylie (Table 9). Home ranges of adjacent groups frequently overlapped—100% overlap reported for some groups—in all three areas (Deag & Crook, 1971, p. 188; Rumsey & Whiten, 1972, p. 562; Menard et al., 1986, p. 37; Mehlman, 1989, p. 282). Of 13 observed visual contacts between neighboring groups, only three (23%) were agonistic (Mehlman & Parkhill, 1988, p. 33; cf. Deag & Crook, 1971, p. 189). Linear day ranges of >2 km have been reported for two Algerian groups (Ménard & Vallet, 1997, ps 2or) Population density in M. sylvanus apparently attains its maximum in relatively undisturbed cedar or mixed cedar/oak habitats, where densi- ties of 19-70 individuals per km* have been reported (Table 10). As expected, degraded habitats, particularly those in the Moroccan Rif, support population densities that are much lower—0.37-4.50 individuals per km? (cf. Mehl- man, 1989, p. 286; Dell’Amico et al., 2006, p. 292). Two semi-natural populations main- tained in France have reached densities of 1,500 and 2,600 individuals per km* (de Turck- heim & Merz, 1984, p. 245). Diet The diet of natural populations of M. sylvanus includes a wide variety of gymnosperms and angiosperms (Fa, 1984d, pp. 348-355) (Ta- ble 11). In the Moroccan Moyen Atlas, 107 species of food plants have been reported FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 15 TasLe 9. Home range area in natural populations of Macaca sylvanus. Sample area Ain Kahla Bou Jirirh Means (weighted) Beni Hessane: Bouhassim Rhomara: Talssemtane Means (weighted) Akfadou Djurdjura: Mount Guessig Djurdjura: Tigounatine Means (weighted) Group size Home range area (ha) References! Morocco: Moyen Atlas 32? (25-39) 15.17 (11.7-18.6) 1 40 25.0 2 34.7 18.4 — Morocco: Rif® 27 1200.0 1 28.8" (12-59) 725.47 (307-901) 3 28.5 804.5 — Algeria: Grande Kabylie 39-45 424 4 >37 39 5 46-73 376 4 >46.2 279.7 ad ' Key to references: /. Fa, 1986d, p. 202. 2. Drucker, 1984, p. 139; Fa, 1986d, p. 202. 3. MehIman, 1989, p. 282. 4. Ménard and Vallet, 1996, pp. 123-124. 5. Ménard et al., 1990, p. 169. 9 ~ . . . ~ Mean value of two groups; extreme values indicated in parentheses. * Cf. Rumsey and Whiten, 1972, p. 562. * Mean value of five groups; extreme values indicated in parentheses. (Drucker, 1984, p. 142; cf. Deag, 1983, p. 573); & Vallet, 1986, pp. 180, 190; 1996, pp. 120, 125; in the Moroccan Rif, 100 species have been cf. Ménard, 1985, pp. 460-463). The 100 species reported (Mehlman, 1988, pp. 472, 477; cf. Fa, exploited as food in the Moroccan Rif constitut- 1983a, p. 84); and in the Algerian Grand ed 51% of 195 species of seed plants identified as Kabylie, 130 species have been reported (Ménard present in that area, and, similarly, the 130 TasBLe 10. Population density in Macaca sylvanus. Sample area Dominant vegetation Survey year Density (individuals/km*) References’ Morocco: Moyen Atlas Afennourir Cedar (protected) 1994-1995 19.3 1 Ain Kahla Mixed cedar/oak 1968-1969 60-70 2 Ain Kahla Mixed cedar/oak 1973-1974 40 3 Azrou vicinity Mixed cedar/oak 1994 28.19 4 Azrou vicinity Mixed cedar/oak 2002 7.26 4 (degraded) Bou Jirirh Cedar (undamaged) 1980-1981 50 5 Cédre Gouraud? Mixed cedar/oak 1994-1995 7.8 1 (degraded) Oued-el-Abid Mixed oak/juniper 1984 ~11 6 Morocco: Rif Anjera: Moussa, Jebel Scrub 1980 1.09 7 Beni Hessane: Bouhassim Mixed oak 1980 0.37 vf Beni Zaid Scrub 1980 0.60 7 Rhomara Various 1980 1.08-4.50 7 Rhomara: Talassemtane Fir 1981-1982 8.3 8 Algeria: Grande Kabylie Akfadou Deciduous oak 1990 12.5 9 Djurdjura: Tigounatine Mixed cedar/oak 1990 28.2 9 ' Key to references: /. Machairas et al., 2003, p. 189; cf. Camperio Ciani and Machairas, 2003, p. 124. 2. Deag, 1977, p. 269; 1984, p. 123. 3. Taub, 1977, p. 110. 4. Camperio Ciani et al., 2005, p. 638. 5. Drucker, 1984, p. 136; Fa, 1986d, p. 202. 6. Fa, 1986b, p. 33. 7. Fa, 1982, p. 62. 8. Mehlman, 1984, p. 175. 9. Ménard and Vallet, 1996, p. 124. * Alternate locality name: Gouroud. 16 FIELDIANA: ZOOLOGY TABLE 11. sylvanus populations. Number of seed-plant Sample area species eaten Number of gymnosperm and angiosperm (seed-plant) species consumed by Macaca References Morocco: Moyen Atlas Deag, 1983, p. 573 Drucker, 1984, p. 142 Morocco: Rif Fa, 1983a, p. 84 Mehlman, 1988, pp. 472, 477 Algeria: Grande Kabylie Ain Kahla 70 Bou Jirirh 107 Djebala region 46 Lakraa, Jebel 100! Akfadou WV Icetcifére 3 It Tigounatine 130° Ménard, 1985, pp. 460-463; Ménard & Vallet, 1996, pp. 120, 125 Menard & Vallet, 1986, p. 190 Ménard & Vallet, 1996, pp. 120, 125 ' Total seed-plant species identified as present in this sample area: 195 (51.3% consumed). * Total seed-plant species identified as present in this sample area: 271 (48.0% consumed). species exploited in the Algerian Grand Kabylie constituted 48% of 271 species identified as present in that area. Barbary macaques also consume fungi, lichens, mosses, and animal prey, (Dea. 1983; p.571; Fa; 1984d;~p. 355; Ménard, 1985, p. 452). Agricultural crops are known to have been raided by Barbary macaques at least since the early 16th century (Leo Africanus, 1896 edition, p. 948; cf. Mehlman, 1988, p. 477). Plant parts that are eaten by M. sylvanus include flowers, fruits, seeds, seedlings, leaves, buds, bark, gum, stems, roots, bulbs, and corms (Fa, 1984d, p. 348; Mehlman, 1988, p. 472; Ménard & Qarro, 1999, p. 130). Because dietary composition varies seasonally (Deag, 1983, p. 571; Mehlman, 1984, p. 192; Ménard, 1985, p. 455; Ménard & Vallet, 1986, p. 177), compar- ative quantitative evaluation of dietary compo- nents requires study periods that extend over at least 12 months. Studies of that duration have indicated that seeds and leaves are the main foods (ca. 60-75%) of Barbary macaques in the Algerian Grande Kabylie (Table 12). Other studies have suggested that fruits could consti- tute a more significant part of the diet in the Moroccan Moyen Atlas (Deag, 1983, p. 574; Drucker, 1984, p. 142; Ménard & Qarro, 1999, p. 130). Animal prey consumed by M. sy/vanus in- cludes snails, earthworms, scorpions, spiders, centipedes, millipedes, grasshoppers, termites, water striders, scale insects, beetles, butterflies, moths (including larvae), ants (including nests), and tadpoles (Fa, 1983a, p. 84; 1984d, p. 355; Mehlman, 1984, p. 193; 1988, p. 476; Ménard & Vallet, 1986, p. 187). Semi-free-ranging monkeys have been observed to pursue and/or catch birds, squirrels, and young rabbits, but never to eat them (Kaumanns, 1978, p. 59; de Turckheim & Merz, 1984, p. 248); mice apparently are ignored. Feeding reportedly occupied an average of 24% and 25% of daytime hours at two localities in the Moroccan Moyen Atlas (Machairas et al., 2003; «~p! 193; “ef. -Fa, 1986c; ap: 216) and> also occupied 24% and 25% of daytime hours at two localities in the Algerian Grande Kabylie (Meé- nard & Vallet, 1997, p. 291). In winter, water has been obtained by eating snow (de Turckheim & Merz, 1984, p. 249; Mehlman, 1988, p. 482). During other times of water shortage, M. sylvanus has stripped the bark of cedar trees in order to obtain retained moisture (Camperio Ciani et al., 2001, p. 263). Predators Under natural conditions, the leopard proba- bly was a major predator of M. sy/vanus when these two species were broadly sympatric (Au- lagnier & Thevenot, 1986, p. 56; cf. Fa, 1986b, p. 31). Large eagles presumably also were and are predators, because Barbary macaques re- spond to eagles’ approach with alarm calls (Mehlman, 1984, p. 191); domestic dogs elicit a similar response, but foxes and jackals do not (cf. Ménard et al., 1986, p. 36). In semi-free-ranging European populations, M. sylvanus gave alarm calls in response to the appearance of snakes, raptorial birds, and dogs FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 17 TABLE 12. Allocation of feeding time to various dietary components by Macaca sylvanus groups in Grande Kabylie, Algeria (Ménard, 1985, p. 454; Ménard & Vallet, 1986, p. 175; cf. Ménard & Vallet, 1996, p. 122). Dietary component Seeds 32.2 Leaves” 213 Lichens 14.2 Invertebrates 10.5 Roots 6.9 Fungi 4.1 Flowers 3.4 Fruits 0.8 Other 0.6 Totals 100.0 Akfadou' (Deciduous oak forest) Feeding time (%) Tigounatine” (Cedar/evergreen oak forest) th GO ON —-~ SONA SEE NAae SADWAN~AIA © S ' Study period: February 1983—September 1984. > Study period: April 1983—March 1985. > Includes herbs. (Paul & Kuester, 1988, p. 200; Fischer & Hammerschmidt, 2002, p. 37). In a European zoo, an infant Barbary macaque was killed by a hawk (Salzert, 1978, p. 39). Intergroup Migration Transfer of males between M. sy/vanus groups apparently is reduced or postponed relative to intergroup transfer in other macaque species (K[ue]ster & Paul, 1984a, p. 70; Paul & Kuester, 1984, p. 372; Taub, 1984b, p. 52; Kuester & Paul, 1986, p. 317; 1997, p. 962; Ménard & Vallet, 1996, p. 118; Kiimmerli & Martin, 2005, p. 1232). At two localities in the Algerian Grande Kabylie, 50% of sexually mature males remained in their natal groups (Ménard & Vallet, 1996, p. 118; cf. Scheffrahn et al., 1993, p. 383), and, in a semi-free-ranging European popula- tion, ca. 67% of sexually mature males remained in their natal groups (Kuester & Paul, 1999, p. 91; cf. 1989, p. 94). In the Algerian groups, the age of migrating males usually was >5 years (78- 100%); in the European groups, the age of migrating males usually was 3-4 years (extremes, 1.75-11.75 years). Migration most frequently occurred during the fall-winter mating season (Ménard & Vallet, 1996, p. 119; Kuester & Paul, 1999, p. 91). Females rarely migrated (Paul & Kuester, 1988, p. 216; Kuester & Paul, 1989, p. 95; Ménard & Vallet, 1993b, p. 110; cf. Ménard et al., 1986, p. 36). 18 Reproduction Seasonality Reported matings in natural populations of M. sylvanus were restricted to the period September—April (wet season), with a mating peak during the period November—January (Table 13). The mating seasons of semi-natural and captive populations were of generally similar duration, but they might extend slightly longer than those in natural populations. In the Salem semi-natural population, onset of the mating period of individual females was related to age and reproductive history (K[ue]ster & Paul, 1984a, p. 71; 1984b, p. 193); older females (>10 years) began mating earlier in the year than younger females, and adult females without young of the previous year began mating earlier than adult females with young of the previous year. Births in natural populations of M. sylvanus were restricted to the period February—Septem- ber, and births in semi-natural and captive populations were restricted to the period Janu- ary—November (Table 13). The timing of birth seasons can vary locally; the observed median birth date at Akfadou (3 May) was significantly earlier than that at nearby Tigounatine (19 May) (Menard & Vallet, 1996, p. 109), and the birth season at Kintzheim apparently occurs earlier than at Rocamadour and Salem (de Turckheim & Merz, 1984, p. 257). FIELDIANA: ZOOLOGY Tasie 13. Seasonal mating’ periods (m) and birth periods (b) reported in Macaca sylvanus populations. Sample area Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul References” Morocco: Moyen Atlas Ain Kahla m m m m m/b m/b b b b 1 Cédre Gouraud* m mm b b b b p Afennourir b m b b Jy, Morocco: Rif Talassemtane m m m m m m/b b b 3} Lakraa, Jebel b b m m m m b b b 4 Algeria: Grande Kabylie Icetifére m4 b b b b 5) Tigounatine m* b b b b =) Akfadou m m m m m b b b b 5 Semi-natural or captive populations U.K.: Gibraltar b m/b m m m m m m/b m/b b b 6 France: m m m m b b b b b i Rocamadour France: m m m m mb m/b m/b_ m/b b b b 8 Kintzheim Germany: Salem m/b m m m m m m/b b b b b 9 Germany: Rheine b b b b b 10 U.K.: Jersey b b b 11 U.S.A.: Nat. Zoo b b m/b b b b b b b TEL ' Mating is defined as ejaculatory copulation. ? Key to references: 7. Deag and Crook, 1971, pp. 183, 189; Deag, 1984, pp. 124-125. 2. Machairas et al., 2003, p. 189. 3. Mehlman, 1984, p. 176; 1986, p. 69; 1989, p. 276. 4. Rumsey and Whiten, 1972, p. 562. 5. Ménard et al., 1985, p. 73; Ménard and Vallet, 1993b, p. 108; 1996, p. 109. 6. Zeuner, 1952, p. 271; MacRoberts and MacRoberts, 1966, p. 300; 1971, p. 39; Burton, 1972, pp. 45, 52; Burton and Sawchuk, 1974, p. 275; Fa, 1984b, p. 279; 1986c, p. 152; Semple, 1998, p. 288. 7. Small, 1990a, p. 86; Hammerschmidt and Ansorge, 1989, p. 79; Ansorge et al., 1992, p. 272; Will and Todt, 1997, p. 66. 8. de Turckheim and Merz, 1984, p. 256; Prud’Homme, 1991, p. 11. 9. Paul and Thommen, 1984a, pp. 4, 6; K[ue]ster and Paul, 1984a, p. 71; de Turckheim and Merz, 1984, p. 256; Paul and Kuester, 1987, p. 120; 1988, p. 207. 10. Salzert, 1978, p. 39. 77. Mallinson, 1979, p. 37. 12. Roberts, 1978, p. 231. > Alternate locality name: Gouroud. 4 Cycling females (but no copulations) observed in other months. Sexual Maturation In natural populations in the Algerian Grand Kabylie, females typically were at least 5 years old before they produced their first infants (18 of 32 females), which indicates that the minimal age for female sexual maturity (capability of preg- nancy) generally was ca. 4.5 years (Ménard et al., 1985, p. 68; Ménard & Vallet, 1996, p. 114); only 1 of 37 females (3%) in this population produced an infant at age 4 years, and none of 43 females produced an infant at age 3 years. Males in the Algerian Grand Kabylie were capable of ejacu- latory copulation at age 5—7 years, although they were not yet full grown (Ménard et al., 1985, p. 68); the same was true of males at age 5—7 years in the Moroccan Moyen Atlas (Deag, 1980, p37): The chronology of sexual maturation in pro- visioned semi-natural populations apparently was generally similar to that in natural popula- tions (Burton, 1972, p. 48; de Turckhem & Merz, 1984, p. 257; Fa, 1984c, pp. 341, 343). However, maturation might have been somewhat acceler- ated in semi-natural populations: Paul and Kuester (1996b, p. 298) reported that, in the semi-natural Salem population, 55 of 157 females (35%) produced their first infants at age 4 years (see above). Reproductive Anatomy Notes Although morphology of the glans penis and uterine cervix provides important taxonomic characters relevant to species relationships in the genus Macaca (Fooden, 2006, p. 27), little information is available concerning these struc- tures in M. sylvanus. Illustrations of the glans penis published by Daubenton (in Buffon & Daubenton, 1766, p. 119, plate 11; slightly FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 19 revised in Buffon & Daubenton, 1833, plate 415, fig. 1) and Hill (1958, p. 654; 1974, p. 207) indicate that the glans in M. sy/vanus is bilobed and relatively broad, as in the silenus group (Fooden, 1975, p. 33). Hill (1974, p. 621) also has provided measurements of two M. sylvanus bacula: length—11.2 mm, 11.5 mm; dorsoven- tral diameter—3.1 mm, 3.5 mm; and transverse diameter—2.2 mm, 2.0 mm. Illustrations of the female reproductive tract have been published by Daubenton (in Buffon & Daubenton, 1766, p. 121, plate 12) and Rouget and Sabatier (1866, plate 8, fig. 2), but, unfortunately, neither of these figures depicts critical characters of the uterine cervix. Female Sexual Cycles During the mating season, M. sy/vanus females exhibit coordinated reproductive cycles of sexual skin swelling, estrus behavior, and menstruation. The sexual skin cycle consists of prominent tumescence and detumescence of two areas of perineal sexual skin—one located around the anus (Fig. 2C) and the other located bilateral to the vulval cleft (Burton, 1972, p. 49; de Turck- heim & Merz, 1984, p. 256; Fa, 1984c, p. 343; K[ue]ster & Paul, 1984a, p. 75). Developmental- ly, these circumanal and lateral vulval swellings are first evident as slight pinkish bulges that appear in females at age ca. 2.5 years. During subsequent mating seasons, the swellings in- crease in area and thickness—and often become bluish-gray—until age ca. 7-15 years, after which the swellings gradually decrease in size, ultimate- ly becoming inconspicuous beyond age ca. 20 years. At the stage of maximal development, the area of the circumanal swelling (ca. 200 cm?) is approximately four times as large as the area of the lateral vulval swellings (Fa, 1984c, p. 345; MOhle et al., 2005, pp. 355, 359). Of 12 females in the Gibraltar M. sylvanus population that were closely monitored during the 2001-2002 mating season, ten exhibited two cycles of tumescence and detumescence, and two exhibited three cycles (Mohle et al., 2005, p. 358; cf. Roberts, 1978, p. 231); tumescence was associated with an increasing fecal estrogen level, and detumescence was associated with an in- creasing fecal progestagen level. Ten of the monitored females became pregnant during one of their cycles, and all ten of these females exhibited a postconception tumescence. The 20 median interval between successive tumescence peaks was ca. 32 days (extremes, 27-44 days). The estrus cycle consists of a period of female proceptive/receptive sexual behavior, marked by gradually increasing copulation rate, abruptly followed by a period of female sexual inactivity (K[ue]ster & Paul, 1984a, pp. 73, 75; Ménard et al., 1985, p. 73; Paul & Kuester, 1987, p. 125; Kuester & Paul, 1989, p. 96; cf. Small, 1990b, p. 272); maximal copulation rate in an estrus cycle is correlated with maximal sexual swelling turgescence. Most females exhibit two estrus cycles per mating season; of 58 mature females in the Salem semi-natural population, 38 (65.5%) had two estrus cycles during the 1982-1983 mating season, 9 (15.5%) had one estrus cycle, and only one female had a maximum of five estrus cycles during this season (cf. Small, 1990b, p. 271; Semple, 1998, p. 288). A female’s first estrus cycle of the season tends to be the longest, including 2-4 weeks (rarely 6 weeks) of sexual activity and ca. 2 weeks of sexual inactivity; subsequent estrus cycles include about one week of sexual activity. Most females probably are fertilized during the first estrus cycle of the season (K[ue]ster & Paul, 1984b, p. 194). During an estrus cycle, several females in the same group (5 of 13 in one group; 6 of 14 in another) were sexually active concurrently (Semple, 1998, p. 288; cf. Ménard et al., 1985, p. 73). The menstrual cycle in M. sylvanus has a median length of ca. 32 days, including a median duration of menstrual bleeding of 4 days (extremes, 1—9 days) (Panouse, 1957, p. 28; Mohle et al., 2005, p. 360). The median interval between the last day of maximal sexual skin swelling and the first day of menstruation was 16 days (extremes, 12-18 days; n = 11) (MOhle et al., 2005, p. 358). Postconception bleeding in M. sylvanus has been reported to occur ca. three weeks after fertilization (K[ue]ster & Paul, 1984a, p. 75; Mohle et al., 2005, p. 361). Consortships During the mating season of M. sy/vanus, pairs of males and estrous females form consort- ships—temporary sexual associations in the course of which copulations occurred (Taub, 1978b, unpaginated; 1980b, p. 292; de Turkheim & Merz, 1984, p. 257; Fa, 1986c, p. 151). The reported duration of consortships varied from <1 minutes to 93 minutes (median = 13 min- utes, m = 297 consortships) in the Moroccan FIELDIANA: ZOOLOGY Moyen Atlas (Taub, 1980b, p. 308) and from ca. 1 minute to 240 minutes (85% less than 8 min- utes) on Gibraltar (Fa, 1986c, p. 153). In Morocco, the formation of most consortships was initiated by females (55%; males, 28%; mutually originated, 17%), and females also were responsible for the termination of most consortships (72%) (Taub, 1980b, pp. 299, 302; cf. Todt & Pohl, 1984, p. 225; Small, 1992, p. 142). The mean rate of exchange of consort- ship partners by females in the Moroccan Moyen Atlas was one per 19.9 minutes (Taub, 1980b, pp. 306, 308; cf. Small, 1992, p. 144; Paul et al., 1993b, p. 498). In the studied group, which included six mature females and 11 mature males, females consorted with a mean of 6.1 males per day (extremes, 3-10 males) (cf. Fa, 1986c, p. 151; Small, 1993, p. 84; Ménard et al., 2001, p. 606). In this group, a single copulation occurred during most consortships (198/278 = 71%); two copulations occurred during nine consortships (9/278 = 3%); and 71 consortships were nonejaculatory (71/278 = 26%) (Taub, 1980b, p. 314). Copulation As in other macaques, a M. sylvanus male typically mounts a M. sylvanus female dorsoven- trally by gripping her trunk and shanks with his hands and feet, respectively (Fig. 8; cf. Fooden & Aimi, 2005, p. 60). This species is a single-mount ejaculator—that is, copulations usually are re- stricted to a single mount—dismount sequence, during which the participating male ejaculates following a brief series of pelvic thrusts (Taub, 1980b, p. 313; 1982, p. 110; Mehlman, 1986, p. 72; Kuester & Paul, 1989, p. 99; 1992, p. 208; van den Bergh, 1991, p. 40; Semple, 1998, p. 288). In the Ain Kahla group, studied in Morocco, the mean (+SD) duration of 205 observed copulations was 8.7 + 1.1 seconds (extremes, 6—14 seconds), and the mean number of pelvic thrusts to ejaculation was 9.1 + 1.8 (extremes, 5—21) (Taub, 1980b, pp. 313, 314; cf. Mehlman, 1986, p. 72; Kuester & Paul, 1989, pp. 99, 100; 1992, p. 208); the mean interval between consortship formation (see above) and onset of copulation was 2.3 minutes (extremes, <1l-16 minutes). Although harassment of copu- lating pairs by other group members was not observed at Ain Kahla (Taub, 1980b, p. 296), 100 incidents of harassment by females were observed in the Salem semi-natural population (Kuester & Paul, 1996, p. 770); of these harass- ment incidents, only 5% resulted in successful displacement of the harassed target. During the peak mating season at three Algerian localities, Ménard et al. (1985, p. 73) observed an average of 1.1—-1.3 copulations per hour. During the concluding phase of a copulation, the female partner utters a characteristic series of loud, low-frequency grunts (Taub, 1980b, p. 296; Todt & Pohl, 1984, p. 225; Paul, 1989, p. 463; van den Bergh, 1991, p. 40; Small, 1992, p. 142; Todt et al., 1995, p. 153; Semple, 1998, p. 288). Males apparently are able to detect differences between copulation calls produced by females at different stages of the estrous cycle (Pohl & Todt, 1984, p. 373; Semple & McComb, 2000, p. 710; Fischer & Hammerschmidt, 2002, p. 40; cf. Pfefferle et al., 2006, abstract no. 579). Male Dominance Rank and Reproductive Success Early studies of the relationship between male dominance rank and reproductive success in M. sylvanus were based on observation of copula- tion frequency (mating success), whereas more recent studies have been based on _ genetic determination of paternity (fertilization success). Of ca. 234 copulations observed in a natural group at Ain Kahla, 76.6% were accumulated by four high-ranking males, and only 23.4% were accumulated by seven low-ranking males (Taub, 1980b, p. 320). Similarly, in the semi-natural population at Salem, mating success was strong- ly dependent on dominance rank (Paul, 1989, p. 467). The results of paternity determination stud- ies—in semi-natural groups—are equivocal. Al- though a positive overall relationship between rank and fertilization success was found in the Salem population (Paul & Kuester, 1996b, p. 302) and in the Rheine Zoo population (Witt et al., 1981, p. 207), no such relationship was found in the Gibraltar Middle Hill population (Kiimmerli & Martin, 2005, p. 1238). Inbreeding/Outbreeding Male emigration and group fission tend to separate maternally related M. sy/vanus males and females and thereby tend to reduce the possibility for inbreeding to occur (Kuester & Paul, 1999, p. 101; Lathuilliere et al., 2004, p. 175; see above, Natural History). Even when M. sylvanus males became sexually mature while FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 21 Fic. 8. Morocco, by G. Fooden. still in their natal group, matrilineal inbreeding apparently was rare (Taub, 1980b, p. 334; Paul & Kuester, 1984, p. 372; cf. Merz, 1976, p. 62); in the Salem semi-natural population, incestuous copulations were observed in only 15 of 371 possible matrilineal male-female pairs (Kuester et al., 1994, p. 1186; cf. Paul & Kuester, 1985, p. 320). Avoidance of inbreeding in the Salem population was inferred to be a proximate effect of close association of maternal relatives during early life (Kuester et al., 1994, p. 1190). Gestation In the Salem semi-natural population, gesta- tion length was measured as the interval between the date of an infant’s birth and the date of its mother’s last copulation during a conceptional estrus (Paul & Kuester, 1987, p. 126; cf. Didier & Rode, 1936, p. 11; MacRoberts & MacRoberts, 1966, p. 302; K[ue]ster & Paul, 1984a, p. 77). Copulation posture in Macaca sylvanus. Photographed 29 September 2006 at Cédre Gouraud, The mean (+SD) gestation period determined for 67 female infants was 165.3 + 4.55 days (extremes, 145-178 days), and that for 56 male infants was 163.2 + 5.38 days (extremes, 145-177 days). The difference between the mean gestation periods of female infants and male infants is statistically significant (P < 0.02, f-test). Parturition Details have been reported concerning one M. sylvanus parturition that was observed in 1987 in the semi-natural Rocamadour population (Ham- merschmidt & Ansorge, 1989, p. 79). The ob- served parturition was one of 24 that occurred in this group—all apparently at night or early in the morning—during the March—June birth season. The prepartum female, already mother of a 1- year-old male, drew the attention of observers at 2125 hours (exact date in 1987 unspecified), when she remained sitting alone on the ground while other group members moved into sleeping FIELDIANA: ZOOLOGY TaBLe 14. Mean annual birth rate (births/reproductive years) in Macaca sylvanus groups. Observation Reproductive Mean annual Sample area period Births years birth rate References” Natural groups Rhomara, Morocco 1980-1983 43 74 0.58 1 Tigounatine, Algeria 1982-1990 99° 173 0.57 2 Akfadou, Algeria 1982-1990 67* 105 0.64 2 Totals — 209 352, 0.59 — Semi-natural groups Queen’s Gate, Gibraltar 1936-1994 130 268 0.49 3 Middle Hill, Gibraltar 1946-1994 129 204 0.63 2; Middle Hill, Gibraltar 1995-2001 ey) 64 0.89 4 Rocamadour, France 1987 26 48 0.54 5 Salem, Germany 1978-1988 730 952 0.77 6 Totals — 1072 1536 0.70 — ' Potential reproductive years of sexually mature females in each group. > Key to references: 7. Mehlman, 1989, p. 277. 2. Ménard and Vallet, 1993b, p. 107; 1996, pp. 107, 109. 3. Fa, 1984b, p. 275; Fa and Lind, 1996, pp. 238, 240; cf. Burton and Sawchuk, 1982, p. 142. 4. Kimmerli and Martin, 2005, p. 1240. 5. Small, 1990c, p. 297. 6. Paul et al., 1993a, p. 110. > Midpoint value in range of uncertainty (97-101) specified in source. * Midpoint value in range of uncertainty (66-68) specified in source. trees ca. 30 m distant; during the next 38 min- utes, the solitary prepartum female manually inspected her perineum and emitted regular, low- amplitude vocalizations. Meanwhile, at 2145, after the female had moved on the ground ca. 15 m toward other group members, a liquid— suspected to be amniotic fluid—was noted on her hindquarters. At 2203, the neonate was rapidly delivered directly into the mother’s hands, and, immediately following its birth, the infant emitted at least 12 screams. At 2211, after the mother had licked the infant for a few minutes, the placenta was expelled, and, by 2215, it was completely consumed by the mother. During this stage, the female and infant were joined by a young juvenile, which might have been the female’s 1-year-old offspring. Finally, by 2219, the mother, carrying her infant, and the accompanying juvenile had climbed into a sleep- ing tree. Birth Rate The mean annual birth rate in M. sylvanus was 0.59 in three natural groups and 0.70 in five semi- natural groups (Table 14), which suggests that births every year are relatively uncommon in females of this species—particularly in natural groups (cf. Deag, 1980, p. 57; 1984, p. 125). Among factors known to affect the birth rate are natural food supply fluctuations (Ménard & Vallet, 1993b, p. 111), age and parity of sexually mature females (Paul & Thommen, 1984b, p. 186; Paul & Kuester, 1988, p. 207; Paul et al., 1993a, pp. 109, 110; Ménard & Vallet, 1996, p. 109); and human disturbance of semi-natural groups (Fa, 1988, p. 77). Neonatal Sex Ratio Of 1040 reported M. sy/vanus neonates, 538 were males and 502 were females (male/female ratio = 1.07) (Table 15); although the overall difference was not statistically significant (P > 0.10, chi-square test), the number of male neonates equaled or exceeded that of female neonates in five of the seven available samples. Neonatal sex ratio was related to maternal rank in the Salem sample (Paul & Kuester, 1990, p. 288): the male/female ratio among infants produced by high-rank mothers (102/74 = 1.38) significantly exceeded the ratio among infants produced by low-rank mothers (86/95 = 0.91; P < 0.05, chi-square test); the ratio among infants produced by mid-rank mothers was intermediate (92/96 = 0.96). Twinning In a composite sample of 874 births reported in semi-natural populations of M. sylvanus, FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 2a Taste 15. Neonatal sex ratio in Macaca sylvanus. Number of births Observation Ratio: : Sample area period Females Males Total males/females References Natural groups Tigounatine, Algeria 1982-1990 53 43 96 0.81 1 Akfadou, Algeria 1982-1990 29 34 63 ad. 1 Subtotals -- 82 77 159 0.94 _ Semi-natural groups Queen’s Gate, 1936-1981 59 62 121 1.05 2 Gibraltar Middle Hill, Gibraltar 1936-1981 52 71 123 1.37 2 Middle Hill, Gibraltar 1995-2001 31 35 66 1.13 3 Rocamadour, France” 1987 13 13 26 1.00 df Salem, Germany~ 1977-1988 265 280 545 1.06 5 Subtotals - 420 461 881 1.10 Totals _ 502 538 1,040 1.07 — ' Key to references: /. Ménard and Vallet, 1996, p. 114. 2. Fa, 1984b, p. 283; 1986c, p. 292; cf. Burton and Sawchuk, 1982, p. 142. 3. Kimmerli and Martin, 2005, p. 1240. 4. Small, 1990c, p. 299. 5. Paul and Kuester, 1990, p. 288. > Cf. de Turckheim and Merz, 1984, Ds2ou. twinning frequency was 0.80% (Table 16). Via- bility of M. sy/vanus twins was relatively low; of the 10 pairs reported, 6 pairs were stillborn, 2.5 pairs died within 8 days after birth, and only 1.5 pairs survived. Geissmann (1990, p. 390) has shown that observed twinning frequency in primates tends to decrease as sample size increases to ca. 1,500 births. This indicates that larger samples will be required in order to provide a more reliable estimate of twinning frequency in M. sylvanus. Stillbirths, Infant Mortality Stillbirth frequency (stillbirths/total births) in four semi-natural populations varied from 0.02 to 0.07 (Table 17). Infant mortality rate (infant deaths/infant births)—reported for varying post- natal periods—varied from 0.23 to >0.80 in four natural populations and from 0.04 to 0.10 in four semi-natural populations (Table 18). The con- spicuously lower infant mortality rate reported in semi-natural populations presumably is primar- ily attributable to artificial provisioning. TABLE 16. Twinning frequency reported in semi-natural populations of Macaca sylvanus. Births Twin births Twinning Sample area Observation period reported reported frequency References’ Queen’s Gate, Gibraltar 1936-1981 130 3° 2.3% 1 Middle Hill, Gibraltar 1946-1981 129 i* 0.78% 1 Rocamadour, France 1986 2 ig — 2 Rocamadour, France 1987 25 l 4.0% 25) Salem, Germany 1978-1985 615 ce 0.49% Dayo Gibraltar/Salem totals — 874° ae 0.80% — ' Key to references: /. Fa, 1984b, p. 276; cf. Burton and Sawchuk, 1974, p. 276; Burton and de Pelham, 1979, p. 106. 2. Schaub, 1988, p. 109. 3. Hammerschmidt and Ansorge, 1989, p. 79; Small, 1990c, p. 299. 4. Paul and Kuester, 1988, p. 207; cf. Paul and Kuester, 1987, p. 120. * Stillborn. * One pair stillborn; one pair died 3-4 days after birth. * One pair stillborn; three of remaining four individuals died 2-8 days after birth. > Rocamadour samples omitted because of small number of births reported. 24 FIELDIANA: ZOOLOGY TABLE 17. Sample area Study period Total births Gibraltar: Queen’s Gate 1936-1981 130 Gibraltar: Middle Hill 1948-1981 129 France: Rocamadour 1987 26 Germany: Salem 1977-1985 615 Stillbirth frequency (stillbirths/total births) in semi-natural populations of Macaca sylvanus. Stillbirths — Stillbirth frequency References’ 9° 0.07 | 6° 0.05 | l 0.04 2 12 0.02 i ' Key to references: /. Fa, 1986c, p. 292; ef. Burton and Sawchuk, 1974, p. 276. 2. Small, 1990c, p. 299. 3. Paul and Kuester, 1988, p. 207. ? Includes three twin stillbirths. > Includes one twin stillbirth. In the Salem semi-natural population, neo- natal mortality was greater in males than in females (Paul & Kuester, 1988, p. 213), and the rate of stillbirths and infant mortality was greater in primiparous females than in multipa- rous females (Paul and Kuester, 1996, p. 301). Weaning Macaca sylvanus infants begin to experiment with solid foods at age ca. 45 days, and, during the next few months, mothers often begin to resist nursing attempts by their infants (Burton, 1972, pp. 38, 39; Rumsey & Whiten, 1972, p. 562; Fa, 1984c, p. 339). Although an infant generally continues to nurse sporadically until age ca. | year—or longer, if its mother does not produce a new infant—an orphan in a natural population was seen to be fully capable of feeding itself at age 6-8 months (Deag, 1980, TABLE 18. Macaca sylvanus. p. 57; K[ue]ster & Paul, 1984a, p. 78; Paul & Thommen, 1984a, p. 12; 1984b, p. 188; Ménard et al., 1985, p. 68). In the Salem semi-natural population, older mothers reportedly wean their infants later than younger mothers (Paul et al., 1993a, p. 117). Carpentier (1931, p. 275) reported that a young captive M. sy/vanus infant (with blackish pelage) was successfully nursed by a female dog. Male—Infant Interactions Interactions between infants and older males apparently are more frequent in M. sy/vanus than in most other species of macaques and baboons (Fa, 1984a, p. 8; Taub, 1985, p. 863; Maestri- pieri, 1998, p. 250; Thierry, 2000, p. 119). Nine- teenth century observers noticed that M. sy/va- nus infants on Gibraltar frequently were carried by older males (Lydekker, 1893-1894, p. 121; Infant mortality rate (infant deaths/infant births) in natural and semi-natural populations of Infant Infant Infant Length of infant Sample area Study period __ births deaths mortality rate death interval References! Natural populations Morocco: Azrou vicinity 1994 ? ? >0.80 6 mo 1 Morocco: Cédre Gouraud” 1995 18 13 0.72 ~6 mo yf Algeria: Tigounatine 1983-1990 78-82 v 0.23-0.27 12 mo 3 Algeria: Akfadou 1983-1990 58-60 ? 0.38—0.40 12 mo 3 Semi-natural populations Gibraltar: Queen’s Gate 1936-1981 130 7 0.05 6 mo 4 Gibraltar: Middle Hill 1948-1981 129 9 0.07 6 mo 4 Gibraltar: Middle Hill 1995-2001 or 2 0.04 3 mo 3] Germany: Salem 1978-1988 706 70 0.10 12 mo 6 ' Key to references: /. Camperio Ciani et al., 1996, p. 71. 2. Machairas et al., 2003, p. 189. 3. Menard and Vallet, 1996, p. 116. 4. Fa, 1984b, p. 291; cf. Burton and Sawchuk, 1982, p. 143. 5. Kiimmerli and Martin, 2005, p. 1240. 6. Paul et al., 1993a, p. 110. * Alternate locality name: Gouroud. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE Forbes, 1894, p. 7; cf. Sanderson, 1957, p. 133), and, subsequently, Stewart (1958, p. 340) re- ported that infants that were carried often were taken from their mothers immediately after birth. Lahiri and Southwick (1966, p. 260) studied two newborn infants in captive groups at the National Zoological Park, Washington, D.C., for 12 weeks and found that the infants consistently were associated with the dominant males in their respective groups for ca. 8% of the observation hours; during these interactions, the infants were held, groomed, or carried by the dominant males. Generally similar interactions between infants and adult males (without grooming, however) were observed by MacRo- berts (1970, p. 93; cf. MacRoberts & MacRo- berts, 1971, p. 46; Burton, 1972, p. 33) during a six-month study conducted on Gibraltar. In addition to dyadic male-infant interactions as described above, Deag and Crook (1971, p. 191; cf. Rumsey & Whiten, 1972, p. 562; Deag, 1980, p. 57), who studied natural M. sylvanus groups for two months in the Moyen Atlas, Morocco, observed triadic interactions involving one infant and two older males. In these interactions, one of the males—typically lower ranking—carried an infant to another male—typically higher ranking—and presented the infant to the higher ranking male. Deag and Crook hypothesized that presence of the infant tended to reduce tension that would otherwise exist between a low-ranking male and a nearby high-ranking male; accordingly, Deag and Crook designated dyadic interactions as ‘‘male care” and triadic interactions as “agonistic buffering.” Evidence concerning possible harm to infant participants in male—infant interactions is equiv- ocal. Although Taub (1980a, p. 189) reported that older males handled infants carefully and that no injuries were sustained by infants in 535 interactions that he observed in the Moyen Atlas, Fa (1986c, p. 75) indicated that three infant deaths on Gibraltar were the result of “kidnapping” by the dominant male, and Paul and Kuester (1988, p. 216) attributed at least nine infant deaths at Salem, Germany, to a similar cause. At Kintzheim, France, males have been observed to carry and otherwise interact with dead infants in approximately the same way that they interact with living infants (Merz, 1978, p. 750). The functional significance of triadic male— infant interactions has been vigorously debated. 26 On one side of this debate, an exploitative agonistic-buffering function has been assigned to these triadic interactions by Deag and Crook (1971, p. 191; cf. Deag, 1980, p. 57; Small, 1993, p. 162; Paul & Kuester, 1996a, p. 36); on the other side, an altruistic kin-investment function (i.e., males caring for their kin) has been proposed by Taub (1975, p. 333; 1978b, unpagi- nated; 1980a, p. 196; 1984b, p. 36; 1990, p. 340; Taub & Redican, 1984, p. 381; cf. Smith & Pfeffer-Smith, 1982, p. 105; Riechelmann et al., 1994, p. 285). Significantly, recent genetic studies have not indicated that male—infant triads preferentially consist of closely related individu- als (Kuester & Paul, 1986, p. 322; 2000, p. 92; Paul et al., 1992, p. 96; 1996, p. 163; Ménard et al., 1992, p. 170; 2001, p. 607). Longevity, Reproductive Senescence, Death Rate The greatest life span reported for M. sy/vanus females is ca. 30 years, in the semi-natural population at Salem (Paul et al., 1993a, p. 113; cf. Burton & Sawchuk, 1974, p. 275; 1982, p. 141), and the greatest life span reported for males is ca. 25 years, also at Salem (Kuester et al., 1995, p. 467, cf. Burton, 1972, p. 54; Burton & Sawchuk, 1974, p. 275). The mean postreproduc- tive life span for females is ca. 6 years (extremes 3— 8 years) in seven individuals at Salem that died after reaching estimated ages of at least 25 years (Paul et al., 1993, p. 113; cf. K[ue]ster & Paul, 1984a, p. 71; Paul & Thommen, 1984b, p. 186; Paul & Kuester, 1988, p. 208; 1996b, p. 307); in this group, females continued to cycle for 1-3 years after ceasing to produce infants. The reported postreproductive life span in the 25- year-old male at Salem (see above) was one year (Kuester et al., 1995, p. 467). The mean annual death rate in three semi- natural populations of M. sylvanus varied from 3.4% to 5.8% (Table 19; cf. Fa, 1984b, p. 294). Population Growth Rate The mean annual population growth rate in two natural Algerian populations of M. sy/vanus varied from 4.8% in a deciduous oak forest at Akfadou to 14.6% in a cedar—oak forest at Tigounatine (Table 20). The growth rate in the Tigounatine population approached that re- ported in the provisioned captive population maintained at Salem, Germany (18.3% growth rate). FIELDIANA: ZOOLOGY TABLE 19. Sample area Study period Rocamadour 1976-1981 5.8 Kintzheim 1976-1981 3.4 Salem 1977-1985 3.6! ' Cf. de Turckheim and Merz, 1984, p. 258. Fossils Circum-Mediterranean macaque fossils—most of which were identified as M. sy/vanus or M. cf. sylvanus—have been reported from ca. 70 localities in a broad area that includes the present North African range of M. sy/vanus and also includes much of Europe (Fig. 9, Appendix V). The European records extend as far north as southern England and southern Germany and extend as far east as Romania and the Caucasus; isolated southeastern Mediterra- nean records also have been reported in Israel, Egypt, and Libya. Contrastingly, only one macaque fossil has been reported south of the present North African range—in central Tunisia. It also is noteworthy that no macaque fossils have been reported from Gibraltar (Szalay & Delson, 1979, pp. 355-356; cf. Imrie, 1798, pp. 198, 201; Busk, 1877; p. 129:- Calderon, 1877, p. 128). The oldest circum-Mediterranean macaque fossils were collected at four localities that probably are of Late Miocene age, ca. 7—5.5 million years ago (Ma). These Late Miocene localities are: Almenara-M, east-central Spain, fossils equivocally identified as M. cf. sylvanus (cf. Kohler et al., 2000, p. 450; Eronen & Rook, Mean annual death rate (%) Mean annual death rate in semi-natural populations of Macaca sylvanus. References de Turckheim & Merz, 1984, p. 258 de Turckheim & Merz, 1984, p. 258 Paul & Kuester, 1988, p. 212 2004, p. 327); Menacer (=Marceau), northern Algeria, fossils identified as Macaca sp.; Sahabi area, northeastern Libya, fossils identified as cf. Macaca; and Wadi Natrun, northeastern Egypt, fossils identified as M. libyca (Appendix V). These circum-Mediterranean macaque fossils apparently are older than the oldest known Asian macaque fossils (Yushe Basin, Hubei, China, probably Early Pliocene (ca. 4.5— 4.0 Ma) (Delson, 1980a, p. 20; 1996, p. 40; pers. comm. | January 2007). Most Pliocene macaque fossil localities in Europe are south of 46°N (Fig. 9); the sole exception is Gundersheim, Germany, which is at 49°35'N. Pleistocene macaque fossils in Europe have been reported as far north as ca. 53°N. The youngest macaque fossils reported in Europe are from Solano del Zamborino (Spain), Grotta degli Orsi Volanti (Italy), Torre in Pietra (Italy), and Kugelsteinhohle (Austria)—all Late Pleisto- cene, probably Eemian interglacial, ca. 0.120 Ma—and from Cova Negra (Spain)—Late Pleistocene, last glacial, probably <0.100 Ma (Appendix V). Most Pleistocene macaque fossils in Europe date from interglacial phases (Kurtén, 1968, p. 59; Delson, 1975b, p. 48; 1980a, p. 19). The above discussion ignores an _ isolated cercopithecine lower third molar, probably of TABLE 20. Demographic growth rate in natural and semi-natural populations of Macaca sylvanus. Demographic factors Sample area Forest type Algeria: Akfadou Deciduous oak Study period 1983-1990 Initial census 33 Terminal census 53 Increase factor 1.614 Mean annual growth rate 4.8%° ' Reference: Ménard and Vallet, 1993b, p. 105. * Reference: Paul and Kuester, 1988, pp. 202, 203. * Three daughter groups: 55 + 30 + 21 individuals. * Includes immigration/emigration balance. > Excludes immigration/emigration balance. Natural populations’ Algeria: Tigounatine Semi-natural population” Germany: Salem Cedar—oak Beech—spruce 1983-1990 1976-1983 38 164 106° 533 79° 3.25 14.6% 18.3% FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE ea e Extant populations © Holocene extinctions ible A Pleistocene fossils a B Pliocene fossils C Miocene fossils : ‘oem (\ \ et 25° 30° 155 20° Fic. 9. Geographic distribution of circum-Mediterranean macaque fossils (cf. Appendix V) compared with geographic distribution of extant populations and Holocene extinct populations of Macaca sylvanus (cf. Fig. 1) Late Miocene age, which was collected in Congo (Zaire), central Africa (Hooijer, 1963, p. 26; 1970, p. 166). Although this tooth was originally identified equivocally as “Cf. Macaca c.q. Mesopithecus spec.,” its generic allocation re- mains problematic (Delson, 1980a, p. 11). Systematics Synonymy Macaca sylvanus (Linnaeus, 1758) [[1Lk8nKocg (=Pithekos)]: Aristotle, 384-322 B.C.E., (1984 edition), p. 798 (see McDermott, 1938, p. 88)—external characters; pre-Linnae- an, unavailable name. Galen, 129-200 C.E. (see McDermott, 1938, p. 97)—anatomy. Babuini: Leo Africanus, 1526 (1896 edition), p. 948—external characters; behavior; distri- 28 bution, Mauritania, Bejaia and Constantine mountains; pre-Linnaean, unavailable name. Simia: Gesner, 1551, p. 957—external charac- ters; behavior; pre-Linnaean, unavailable name. Brisson, 1756, p. 188—habitat, Africa. Simia cynocephalus alter: Alpini, ca. 1609 (1979 edition), p. 241, pl. 16 (see Linnaeus, 1758, p. 25)—pre-Linnaean, unavailable name. Cercopithecq: Jonston, 1650 (1657 edition), plate 59, fig. (lower right corner) (see Linnaeus, 1758, p. 25)—pre-Linnaean, unavailable name. Simia ecaudata, clunibus tuberosis: Linnaeus, 1748, p. 3—pre-Linnaean (=pre—1758), un- available name. Simia Cynocephala: Brisson, 1756, p. 191—pre- Linnaean (=pre—1758), unavailable name. Le Pitheque: Buffon, 1766, p. 84—vernacular, unavailable name. Buffon, 1789, pp. 30-36— external characters; behavior. Le Magot: Buffon, 1766, p. 109—vernacular, unavailable name; external characters; behav- FIELDIANA: ZOOLOGY ior; distribution, Tartary, Arabia, Ethiopia, Malabar, Barbary, Mauritania, Cape of Good Hope. F. Cuvier, 1819, pp. 1-4—behavior, measurements; distribution, Barbary, Egypt, southern Spain. Pygmy Ape: Pennant, 1771, p. 98—vernacular, unavailable name. Barbary Ape: Pennant, 1771, p. 100—vernacu- lar, unavailable name. Le Petit Cynocéphale: Buffon, 1789, p. 37— vernacular, unavailable name; compared with Le Magot. Daudin in Lacépéde and Daudin, 1802, p. 149—recognized as a distinct species of Macaca, but not named. [Simia] Sylvanus Linnaeus, 1758, p. 25. Type localities “Africa, Zeylonica.”’ Miller, 1773, p. 120—external characters. Latreille, 1804a, p. 178; 1804b, p. 294—taxonomic relationship to S. inuus uncertain. G. Cuvier, 1817, p. 108—external characters; taxonomy; distri- bution, “Barbarie” (=Barbary Coast). Virey, 1819, p. 275—taxonomy; distribution, juxta- Mediterranean Africa, Gibraltar. Simia Silvanus: Schreber, 1774, p. 68—incorrect subsequent spelling of specific name. [Simia] siluanus Erxleben, 1777, p. 11—unjustified emendation of specific name; synonymy; distri- bution, Africa, India, Jamaica. Macaca sylvanus: Daudin in Lacépéde & Dau- din, 1802, p. 149—new combination; listed as “Variété A” of Macaca inuus. Magus sylvanus: Lesson, 1827, p. 43—new com- bination; taxonomy; distribution, Barbary, Egypt, introduced on Gibraltar. Inuus sylvanus: Jardine, 1833, pp. 149, 214—new combination; taxonomy; behavior, external characters; distribution, warmer parts of Africa and India. Pithecus [(Pithecus)| sylvanus: Dahlbom, 1856, p. 121—new combination; taxonomy; external characters; distribution, North Africa, Gibral- tar. Macacus sylvanus: Schlegel, 1876, p. 115—in- correct subsequent spelling of generic name; external characters; distribution, Morocco, Gibraltar, Algeria. [Macacus (Pithecus)] sylvanus: Trouessart, 1882 (18787), p. 126—new combination; distribu- tion, Algeria, Morocco, Gibraltar. [((Simia or) Macaca] sylvanus: Thomas, 1911, pp. 121, 125—nomenclature; designated type species of Simia by ““Linnaean tautonymy” (cf. International Code of Zoological Nomencla- ture, 1999, Art. 68.5). Macaca _ sylvana: Flower, 1931, p. 155—new combination; unwarranted change of gender ending of specific name; longevity in captivity. [Macaca] sylvanus—group: Miller, 1933, p. 5— new rank; external characters. [| Simia] Inuus Linnaeus, 1766, p. 35. Type locality unknown; name based on “‘Simius cynocepha- lus alter” of Alpini (ca. 1609, p. 241, pl. 16) and Simia cynocephala of Brisson (1756, p. 191); external characters said to be similar to those of S. sy/vanus. Miller, 1773, p. 121— distribution, Africa. Blumenbach, 1788, p. 66—said to be difficult to distinguish from S. sy/vanus; distribution identical to that of S. sylvanus. Audebert, 1797-1798, p. 1— possibly a synonym of S. sy/vanus. Desmarest, 1804, p. 8—species allocated to genus Cynoce- phalus. Macaca inuus: Lacépéde, 1799, p. 4—new com- bination. Stiles and Orleman, 1927, p. 24— validly designated as type species of genus by monotypy; now regarded as a synonym of M. sylvanus (Linnaeus, 1758). Pithecus inuus: E. Geoffroy, 1803, p. 26—new combination; distribution, Africa. Cyn{ocephalus] inuus: Latreille, 1804b, p. 293— new combination; said to be a degenerate “variété” of Le Magot. [M]acacus inuus: Desmarest, 1820, p. 67—in- correct subsequent spelling of generic name. Macacus innus: Desmarest, 1823, p. 470—incor- rect subsequent spelling of generic and specific names. S[imia] Plithecus] inuus: Griffith, 1827, p. 19— new combination. Simia Innuus: Wagler, 1830, p. 5—incorrect subsequent spelling of specific name. M{acacus] Innuus: Gulliver, 1842, p. 68—incor- rect subsequent spelling of generic and specific names. [Macacus (Pithecus)| innuus: Trouessart, 1897, p. 26—new combination; incorrect subsequent spelling of generic and specific names. [Macacus (Innuus)| innuus: Trouessart, 1904, p. 17—new combination; incorrect subsequent spelling of generic and specific names. [Cynocephalus Inuus, variety] %. natatorius Fischer, 1813, p. 539. Type localities ‘Arabia, India, Malabaria, Bucharia?”’; new variety, based on specimen with membranes between pedal digits. Simia Pithecia Link, 1795, p. 60 (not Linnaeus, 1766, p.40)—invalid junior homonym§ of FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 29 [Simia] Pithecia Linnaeus, 1766, p. 40 (= Pithe- cia pithecia, white-faced saki). Simia Pithecus Schreber, [1800], suppl. plate 4B. Type locality unknown; name _ incorrectly attributed to Bufflon] (cf. Wagner, 1839 [1840], p. 149). Inuus pithecus: Lesson, 1840, p. 99—new combi- nation; distribution, Barbary coast, Gibraltar, Alexandria, Cairo, Morocco. I. Geoffroy, 1851, p. 31—said to be indigenous on Gibral- tar. S[imia] Anomela Link, 1795, p. 60. Type locality unknown; name based on Le Petit Cynoce- phale of Buffon, 1789, p. 37. Simia Cynopis Goldfuss, 1809, p. 53. Type locality unknown; name based on Le Petit Cynocephale of Buffon, 1789, p. 37. L{asiopyga]| ecaudis Mlliger, 1811, p. 68—Type locality unknown; name based on Le Petit Cynocephale: Buffon, 1789, p. 37. Inuus ecaudatus E. Geoffroy, 1812, p. 100— Type locality “toute la partie d’Afrique qui cotoie la Méditeranée, et en Europe, Gibral- tar’; external characters; pelage color varia- tion. Innuus ecaudatus: Schinz, 1821, p. 112—new combination; incorrect subsequent spelling of generic name. M{acacus] ecaudatus: Schinz, 1844, p. 60—new combination. I[nuus (Inuus)| ecaudatus: Wagner, 1855, p. 59— new combination. Inuus macacus Rippell, 1835, p. 8. Type locality “ganzen Kiste der Barbarie bis nach Mar- occo”’; name based on captives transported to Egypt. Macacus magots Smith, 1842, p. 103. Type locality “Barbary”; name based on the Bar- bary Ape, probably of Pennant (1771, p. 100). P{ithecus| Pygmaeus Reichenbach, 1862, p. 145. Type locality unknown; name based on the Pygmy Ape of Pennant (1771, p. 98) and a menagerie animal, both of unknown origin; taxonomic comparison. Type Series Although no type specimen was specified in Linnaeus’ (1758, p. 25) original description of Simia_ sylvanus, four bibliographic references were cited. Accordingly, the specimens discussed or figured in these references are syntypes that collectively constitute the type series of Simia sylvanus (International Code of Zoological 30 Nomenclature [ICZN], 1999, Art. 72.4.1). The four cited references are: (1) Gesner, 1551, p. 957—includes figure of tailless monkey; (2) Alpini, ca. 1609, pp. 241-242, plates 15, 16— includes figures of captives transported from Alexandria to Venice; (3) Jonston, 1650 (1657 ed.), pl. 59, fig. (lower right corner)—unnum- bered figure of tailless monkey labeled “‘Cerco- pithecq”’; and (4) Linnaeus, 1748, p. 3, plate 1, fig. 1 (this publication includes another citation of Alpini, ca. 1609, p. 241)—possibly includes ambiguously captioned figure of skull. None of the syntypes is known to have been preserved. At present, designation of a lectotype from among the available syntypes would serve no useful taxonomic purpose (cf. ICZN, 1999, Art. 7403). Type Locality Originally given as “Africa, Zeylona”; cor- rected to “Africa” by Elliot (1913, p. 174); restricted to “Barbary Coast” by Allen (1939, p. 156). Distribution Disjunct areas in the Atlas and Rif Mountains of Morocco and Algeria, extending from ca. 31°15'N, 7°45'W to ca. 36°45’N, 5°20’E (Fig. 1). Diagnosis Prime dorsal pelage color in adults strongly variegated, buffy to reddish mixed with blackish (Fig. 2C). Mean head and body length 556.8 mm in four adult females and 634.3 mm in three adult males. Tail reduced to an inconspicuous fleshy vestige (4-22 mm long) or completely absent. Mean greatest length of skull 118.7 mm in four adult females and 135.5 in five adult males. Remarks Linnaeus (1758, p. 25) originally allocated the Barbary macaque to the genus Simia, but this generic name was subsequently suppressed by the International Commission on Zoological No- menclature (1929, p. 26; cf. Stiles & Orleman, 1927, p. 1). The next oldest available generic name applicable to the Barbary macaque is FIELDIANA: ZOOLOGY TABLE 21. Approximate age! 13-10 Ma 7-6 Ma 6-5 Ma 0.100—0.020 Ma <0.010 Ma ' Ma = million years ago. Subsequent radiation of Macaca in Asia. Macaca Lacépéde, 1799, p. 4. The specific name sylvanus (Latin, woodland god) is a noun stand- ing in apposition to the generic name and therefore is not required to agree in gender with the generic name (ICZN, 1999, Art. 34.2.1). Because available information concerning geo- graphic variation in M. sylvanus is deficient (see above), recognition of subspecies in this species is currently unwarranted. Specimens Examined Total 103: skins and skulls, 33; skins only, 37; skulls or skeletons only, 33 (see Appendix I). Evolution and Dispersal The most comprehensive studies of the evolu- tionary history of M. sylvanus and its relatives have been published by E. Delson (1973, pp. 140, 667, 685; 1975a, p. 207; 1975b, p. 44; 1980a, p. 11; 2000, p. 170; Szalay & Delson, 1979, p. 354). These publications provide the primary basis for the present discussion (Table 21). Although only one species of macaque—M. sylvanus—inhabits Africa, and ca. 20 species of macaques inhabit Asia, fossil evidence indicates that the genus Macaca originated in Africa and that M. sylvanus is a geographic relict of that origin (Simons, 1970, p. 109; Delson, 1973, p. 689; 1980a, p. 23; Szalay & Delson, 1979, p. 334; Stewart & Disotell, 1998, p. R585; Groves, 2001, p. 221). The tribe Papionini which includes macaques (subtribe Macacina) and baboons, mandrills, mangabeys, and geladas (subtribe Papionina)—probably originated in Africa ca. 13-10 million years ago (Ma) (Raaum et al., 2005, p. 250; cf. Szalay & Delson, 1979, p. 327; Delson, 1980a, p. 24; 2000, pp. 168, 170). Through much of the history of these subtribes, Evolution and dispersal of circum-Mediterranean macaques: hypothetical chronology. Evolutionary event Origin of Papionini (in Africa) Origin of Macaca (in northern Africa) Dispersal of Macaca from Africa to Eurasia” Extinction of Macaca in Europe Anthropogenic reduction of North African habitats of Macaca their distributions apparently have been allopat- ric, with macaques inhabiting northern Africa and baboons, mandrills, mangabeys, and geladas inhabiting central and southern Africa; an inter- subtribal barrier in the vicinity of the present Sahara Desert appears to have been operative. The earliest known macaque or near-macaque fossils date from the Late Miocene (ca. 7-6 Ma) in Algeria, Libya, and Egypt (see above, Fossils). Between ca. 6—5.3 Ma, the Mediterranean Sea became desiccated as a result of tectonic move- ments isolating it from the Atlantic Ocean (Loget et al., 2005, p. 414); Delson (1975b, p. 40) has suggested that this might have facilitated dis- persal of ancestral macaques from Africa to Eurasia. The earliest known macaque fossils in Europe (Spain) date from ca. 5.5 Ma, and the earliest macaque fossils in Asia date from ca. 4.5-4.0 Ma in China (see above, Fossils) and from ca. 3 Ma in India (Delson, 1980a, p. 20). DNA evidence supports a sister-group re- lationship between M. sylvanus and all species of Asian macaques (see above, DNA Variation). Judging from tail length variation in extant Asian macaques (Fooden, 2006, p. 27), the ancestral macaque stock that dispersed from Africa to Eurasia probably was long-tailed; if so, this stock probably would not be regarded as conspecific with extant M. sylvanus. After dispersal of macaques from Africa to Eurasia, tail reduction in North African and Eurasian macaques apparently evolved independently at least four times—probably as an adaptation to cool climate (cf. Fooden, 1988, p. 4; 2006, p. 3). Macaques survived in Italy—and perhaps Austria—at least until the Eemian interglacial interval (ca 0.120 Ma), and they survived in Spain at least until the beginning of the last glaciation (<0.100 Ma) (see above, Fossils). During cyclical glacial advances, the geographic distribution of macaques in Europe probably was restricted to more temperate southern areas FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE at (Kurtén, 1968, p. 59; Delson, 1980a, p. 19); macaques might have finally become extinct in Europe as a result of inability to tolerate the most severe stage of the last glaciation (ca. 0.020 Ma). During the Holocene (<0.010 Ma), the population of M. sylvanus in North Africa has been reduced as a result of anthropogenic habitat modification (Mehlman, 1984, p. 182; Thirgood, 1984, p. 36; Camperio Ciani et al., 2005, p. 637). Acknowledgments For access to specimens and generous assis- tance, I am deeply grateful to officials and staff members of institutions cited above (see Materi- als and Methods). Valuable supplementary help and advice were provided by R. Banasiak (FMNH), D. Brandon-Jones (Richmond, Surrey), G. Fooden (Los Angeles), S. B. McLaren (CMNH), M. Mouna (isp), C. L. 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Darwin- ian fitmess in a multimale group of Barbary macaques (Macaca sylvana Linnaeus, 1758): Dom- inance reversals and male reproductive success. Folia Primatologica, 36: 201-211. Zeuner, F. E. 1952. Monkeys in Europe, past and present. Oryx, 1: 265-273. ZIEGLER, T., C. ABEGG, E. MEUAARD, D. PERWITASARI- FARAJALLAH., L. WALTER, J. K. HopGes, AND C. Roos. 2007. Molecular phylogeny and evolutionary history of Southeast Asian macaques forming the M. silenus group. Molecular Phylogenetics and Evolution, 42: 807-816. Appendix I. Specimens Examined (Details), Total 103 SKINS AND SKULLS, 33 ALGERIA, 3. Region unknown (3): BM(NH) 1939.1048, irsN IG 16871, NHMBe 1268. MOROCCO, 8. Central Moyen Atlas (5): Azrou—BM(NH) 1919.8.19.1; Azrou, 5 km S— USNM 476786 (external measurements from field catalog), 476788; Azrou, 18 km S—uvusNm 476789: Khenifra, 22 km E—vusnm 476790 (tail measurement from field catalog). Region un- known (3): IRSN IG 16696, MNHN CG 1831/835, USNM 196984 (locality ““Herrata,” not located). U.K., 5. Gibraltar (5): FMNH 47398, 47409, 47410, UsNM 255979, zmuz 11600. Country unknown, 17. AMNH 17408, 17409, 19014, 70269 (mounted skin, not seen), IRSN IG 16696, IG 16701/7108, IG 16701/7825, IG 17058/7586, IG _ 17058/8587, IG 9295, MCZ 15296, MNHN CG 1939/173, CG 1939/ 1118, NMs 1621, RMNH 4152, zmuz 11601, zsBs 1959/4. SKINS ONLY, 37 42 ALGERIA, 2. Skikda (1): Stora—smcnu) 49b (skull in skin). Region unknown (1): MNHN 283 (skull in skin). MOROCCO, 7. Central Moyen Atlas (4): Ain Leuh—isr 123.001 (listed as tsk 123.002 by Aulagnier & Thevenot, 1986, p. 154); Azrou— isk 123.002 (mounted specimen, on exhibit; listed as 1sk 123.001 by Aulagnier & Thevenot, 1986, p. 154), 132.007; Azrou, 18 km S—usnm 478329 (in fluid; external measurements from _ field catalog). Region unknown (3): tsr_ 100.037, RMNH 5-e, 6-f. U.K., 3. Gibraltar (3): BM(NH) 1939.55], 1939.1049, UsNM 258206 (in fluid). Country unknown, 25. AMNH 6316/7951, BM(NH) 1842.12.29.1, 1911.11.11, unnumbered (old collection; skull in skin), FMNH 60716 (in fluid), irsN IG 10743, IG 9323 MB, stand no. 40 (mounted specimen), stand no. 80 (mounted specimen), MNHN 55a (skull in skin), 55b (skull in skin), 286, 287, 290, CG 1930/72, NHMBa 27, NMS 16837, RMNH l—a, 2-f, 3 (skull in skin), 4d, 4153, usnm 200951, 502460 (in fluid), zsps (unnumbered, mounted specimen). SKULLS OR SKELETONS ONLY, 33. MOROCCO, 13 Central Moyen Atlas (9): Azrou, 18 km S—cMNH 45290, UsNM 476780, 476781, 476782, 476783, 476784, 476785, 476787 (external measurements from field catalog for all seven USNM specimens from this locality); Kheni- fra, 22 km E—usnm 476791. Region unknown (4): isk 500 031, 500 032 (probably listed as Isr 100.078 by Aulagnier & Thevenot, 1986, p. 154), 500.03[?] (probably listed as sr 100.072 by Aulagnier & Thevenot, 1986, p. 154), zsBs 1961/ 188. U.K., 1. Gibraltar (1): irsN IG 17073. Country unknown, 19. AMNH 19178, 2060/ 5484, irsn IG 9071, IG 9165/4218, IG 9165/4219, IG 9295, mcz 7098, Nms 1551, 1552, RMNH 7-a, 9-—c, USNM 543429, zmB 8264, 8271, 16492, zmuz 11602, 11742 (skull size indicates that this specimen is a female), zsss unnumbered female, unnumbered male. Appendix II. Gazetteer of Macaca sylvanus Localities Locality names listed as primary entries in this gazetteer preferentially are official names ap- FIELDIANA: ZOOLOGY proved in gazetteers published by the U.S. Board on Geographic Names—USBGN: Algeria, 1972 and USBGN: Morocco, 1970. (Note that the Arabic word for “mountain” is consistently spelled ‘‘djebel” in USBGN: Algeria and “‘jebel” in USBGN: Morocco.) Supplementary refer- ences that have been consulted are: Kartografiai Vallalat, Algeria map, 1991; International Travel Maps, Morocco map, 2000; and Michelin, Morocco map, no date. Localities of M. sy/vanus that were not found in USBGN gazetteers or in supplementary references are spelled here as in the original sources. Secondary entries, with cross references to corresponding primary en- tries, indicate variant spellings or alternate locality names that occur in published literature. For each primary entry in this gazetteer, the sequence of information presented is as follows: 1. Locality name. 2. Elevation, if reported by collector or observer. 3. Name of region (in italics). 4. Name of country (in capital letters). 5. Coordinates of locality (principal source: USBGN gazetteers). 6. Date of collection or observation. 7. Name of collector or observer. 8. Bibliographic reference (in parentheses), if any. 9. Abbreviated name of museum (see above, Materials and Methods) where specimens, if any, are preserved. 10. Number of specimens avail- able, if any (with indication of part preserved, if skin and skull are not both present). 11. Locality number (italicized), as indicated in distribution map (Fig 1). Abid, Oued el, gorge, Bou Tferda vicinity; Southern Moyen Atlas, MOROCCO; ca. 32°24'N, 5°40’W; observed 20-24 Nov. 1984 by J. E. Fa (1986b, p. 32). 14. Adelma, Oued, 400m; Rif, MOROCCO; 35°13'N, 5°03’W; observed Jul.Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 87). 34. Adelmane, Oued. See Adelma, Oued. Adendoun, Djebel, SSW of Jijel, 800-1200 m; Petite Kabylie, ALGERIA; ca. 36°40'N, 5°35'E; observed 1973-1975 by D. M. Taub (1977. p. A22). 56. Afennourir; Central Moyen Atlas, MOROCCO; 33°16’N, 5°16’W; observed Oct. 1994-Sep. 1995 by I. Machairas (Machairas et al., 2003, p. 186). 22. Agfadou. See Akfadou, Forét de. Aguelmame Aziga. See Azigza, Aguelmane. Ahfa-Timesh, N of Oued Laou; Rif, MOROC- COk cal 4a6°2ANe S2h7'W. sebserved be: forel976 by F. Alvarez and F. Hiraldo (1975, p- 253;°ch Fay, 1983a, piy79)a 32: Ain Kahla, 2010 m; Central Moyen Atlas, MO- ROCCO; 33°15'N, 5°13’W; observed Jul.—Aug. 1968 by J. M. Deag and J. H. Crook (1971, p. 186). Observed Jun. 1973—Dec. 1974 by D. M. Taub (1977, pp. 110, 116). 22. Ain Leuh, 1200-1450 m and 1750 m; Central Moyen Atlas, MOROCCO; 33°17'N, 5°23'W; collected 20 Dec. 1923 by Prince L. Murat (cf. Aulagnier & Thevenot, 1986, p. 154); Isr, 1 (skin only). Reported before 1932 by L. Joleaud (1933, p. 852). Observed 1973-1975 by! DeM> Taub. (1977) p: 116;- cis Fa et-al., 1984, p. 91). 22. Ait Mehammed, S of; Southern Moyen Atlas, MOROCCO; ca. 31°53'N, 6°28’W; observed 1973-1975 by D. M. Taub (1977, p. 119; cf. Fa et al., 1984, p. 97). 4. Ait Mgild. See Sidi Mguid. Ait M’Hammed. See Ait Mehammed. Ait Oubane, Forét d’, Djebel Djurdjura, 1750- 2300 m; Grande Kabylie, ALGERIA; 36°28'N, 4°17'E; observed 1982-1983 by N. Ménard (in Fa et al., 1984, p. 103). 5J. Ait Sokhmane forest, right bank of Oued El Abid; Southern Moyen Atlas, MOROCCO; ca. 32°15’N, 5°50’W; reported in 1931 by L. Joleaud (1933, p. 852). 9. Ait Sokmane. See Ait Sokhmane. Ajdir, Plateau d’, 1600-2000 m; Central Moyen Atlas, MOROCCO; 32°56'N, 5°25'W; ob- served 1973-1975 by D. M. Taub (1977, p> 116; cf. Fa etial., 1984, p: 91). 73: Akfadou, Forét de, 500-1600 m and 800—1300 m; Grande Kabylie, ALGERIA; 36°43'N, 4°33’E; observed 1973-1975 by D. M. Taub (1977, pp? 118, 122: rch) Fat-et ak, 1984, 7p5 104). Observed 1983-1984 by N. Menard (1985, p. 452). Blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). Tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). 52. Akra, Chabet El, N of Kherrata, 1500 m; Petite Kabylie, ALGERIA; 36°32'N, 5°18’E; re- ported in 1931 by L. Joleaud (1933, p. 853). Observed 1973-1975 by D. M. Taub (1977, p. 122; cf. Fa, et al., 1984, p. 105). Tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). 54. Al Hoceima, Rif; MOROCCO; 35°15’N, 3°56’W; reported present in 1920s (Taub, 1978a, p. 249). Reported locally extinct before 1979 by D. M. Taub (1978a, p. 249). 4/. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 43 Ali ou Zrou, Jebel, 1600-2000 m; Central Moyen Atlas MOROCCO; 32°55'’N, 5°25’W; ob- served 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, p. 91). /3. Andjera. See Moussa, Jebel. Anjra massif. See Moussa, Jebel. Annaba. See Edough, Djebel. Annocer, ca. 10 km SW of Aguelmane Azigza; Central Moyen Atlas, MOROCCO; 32°54'N, 5°30'W: observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 94). /3. Aqfadou. See Akfadou, Foret de. Arhbala, N of; Southern Moyen Atlas, MOROC- CO; 32°29'N, 5°39'W; observed 1973-1975 by D. M. Taub (1977, p. 119; cf. Fa et al., 1984, p. 97). 14. Arhbalou N’Irkraouen, above Lake Miami, 1600-2000 m: Central Moyen Atlas, MOROC- CO; ca. 32°54’'N, 5°22'W; observed 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, pp. 91, 94). /3. Ari Bou Iblan. See Bou Iblane, Jebel. Asker, Zaouia, Taguelft region; Southern Moyen Atlas, MOROCCO; 32°10'N, 6°10'W; ob- served 1973-1975 by D. M. Taub (1977, p. 119). 8. Asru. See Azrou. Assaka N’Ouam, NE of Bekrit; Central Moyen Atlas, MOROCCO; 33°06’N, 5°10'W; re- ported before 1978 by Mr. Arnoud (Deag, 1977, p. 276). 20. Assaka-n-Quam. See Assaka N’Ouam. Asserdoun, Zaian circle; Central Moyen Atlas, MOROCCO; 32°41'N, 5°46’W; infant cap- tured 30 Apr. 1929 by local residents (Carpen- tier, 1931, p. 275). JJ. Azigza, Aguelmane, 1600-2000 m; Central Moyen Atlas, MOROCCO; 32°58'N, 5°26'W; collected May 1950 by A. Kalfleche (Aulagnier & Thevenot, 1986, p. 154); Isr 150.011 (not seen); the M. sy/vanus specimen collected at this locality is now missing (searched for on 25 Sep. 2006): the monkey currently tagged with this number has a tail that is longer than the extended hind leg. Observed 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, p. 91). 13. Azrar, Taguelft region; Southern Moyen Atlas, MOROCCO; 32°18’N, 6°00'W; observed 1973-1975 by D. M. Taub (1977, p. 119). Population extant 1982-1984 (Fa et al., 1984, p. 96). 9. Azrou, 1600-2200 m and 1890 m; Central Moyen Atlas, MOROCCO; 33°26'N, 5°13'W; 44 collected 15 Jun. 1919 by H. Lynes (cf. Napier 1981, p. 6); BM(NH), 1. Collected 19 Dec. 1923 by Prince L. Murat (cf. Aulagnier & Thevenot, 1986, p. 154); isk 1 (skin only). Collected in 1932 by J. Surcouf (cf. Aulagnier & Thevenot, 1986, p. 154); isk, | (skin only). Observed 1973-1975 by D. M. Taub (1977, p. 117; ef. Fa et al., 1984, p. 91). 23. Azrou, 5 km S; Central Moyen Atlas, MOROC- CO; 33°24’N, 5°13’W; collected 21 Jun. 1970 by M. G. Hearst; usNM, 1. Collected 21 Jun. 1970 by L. W. Robbins; usnm, 1. 23. Azrou, 18 km S; Central Moyen Atlas, MO- ROCCO; 33°16'N, 5°12'W; collected 22 Jun. 1970 by M. G. Hearst (cf. McLaren et al., 1984, p. 513); cmnu, | (skull only), UsNM, 4 (skulls only). Collected 22 Jun. 1970 by L. W. Robbins; usnm, 5 (1 in fluid, 1 skeleton only, 2 skulls only, | skin and skeleton). 22. Azrou, above, 1520-1700 m (5000-5600 ft); Central Moyen Atlas, MOROCCO; 33°25'N, 5°10'W; observed 2 May-17 Jul. 1919 by H. Lynes (1920, p. 273, pl. 12). 23. Azrou Forest; Central Moyen Atlas, MOROC- CO; ca. 33°26'N, 5°13’W; tissue samples collected before 2006 by L. Modolo et al. (2005; .p.:7393): 23: Azrou region, Central Moyen Atlas, MOROC- CO; ca. 33°26'N, 5°13’W; tissue samples collected from captives before 2006 by L. Modolo et al. (2005, p. 7393; cf. de Turckheim & Merz, 1984, p. 242). 23. Azzaba, W of; Azzaba, ALGERIA; ca. 36°44'N, 7°06'E; unconfirmed local report in 1914 (Joleaud, 1933, p. 853), beyond modern east- ern limit of distribution (Fa et al., 1984, p. 107). 6/. Bab Barred. See Tisirene, Jebel. Bab Besen. See Besene, Bab. Bab Bessen. See Besene, Bab. Babor, Djebel, Kherrata gorge, 1500 m; Petite Kabylie, ALGERIA; 36°30'N, 5°28’E; re- ported in 1925 by L. Joleaud (1926, p. 129; 1933, p. 853). Observed 1973-1975 by D. M. Taub (1977, p. 122; cf. Fa et al., 1984, p. 105). 53: Babors. See Babor. Bab Taza. See Lakraa, Jebel. Bejaia. See Pic des Singes National Park. Bekrit. See Assaka N’Ouam. Beni Ghobri. See Beni Rhobri. Beni Hosmar. See Buzeitune, Jebel; Sidichmim. Beni Hozmar. See Zarka cascade. Beni M’hamed. See Beni Mohammed. FIELDIANA: ZOOLOGY Beni Mohammed, Rhomara tribal area, 1300 m:; Rif, MOROCCO; 35°09'N, 5°07’W; reported Jul—Sep. 1980 by J. E. Fa (1982, p. 53; cf. Fa et al., 1984, p. 87; Mehlman, 1984, p. 168). 34. Beni Rhobri, Forét des, 500-1600 m; Grande Kabylie, ALGERIA; 36°44'N, 4°28’E; Ob- served 1973-1975 by D. M. Taub (1977, p. 122; cf. Fa et al., 1984, p. 104). 52. Beni Touzin. See Beni Touzine. Beni Touzine vicinity, between Metalsa and Ourighel; Rif, MOROCCO; ca. 35°00’'N, 3°42'W; reported present before 1932 by L. Joleaud (1933, p. 852). Reported present before 1962 and locally extinct in 1973-1975 by D. M. Taub (1977, p. 113; 1984a, p. 73). 42. Beni Ziat. See Targa. Besene, Bab, Rif, MOROCCO; ca. 35°00'N, 4°45'W; reported present in mid-1960s and locally extinct subsequently (J. Chitty in Taub, ITT, , 1133-1984a: pp. 73), 37. Blida. See Chiffa, Gorges de la; Chrea. Bone. See Edough, Djebel. Bor) Mira; Petite Kabylie, ALGERIA; ca. 36°34'N, 5°18’E; reported by forestry officials as locally extinct before 1962 (D. M. Taub, 19 jp. 22: ef: Faet al.,. 1984. p. 105). 54. Bouak, Cap and Cap Carbon, between; Grande Kabylie, ALGERIA; ca. 36°47'N, 5°06’E; reported in 1931 by L. Joleaud (1933, p. 853; etmba et abs 1984, p. 105). 53. Bou Cédre, between Azigza Aguelmame and Senoual, 1800 m; Central Moyen Atlas, MO- ROCCO; ca. 32°58’N, 5°20'W; observed 1973-1975 by D. M. Taub (1977, p. 116; Fa etal., 1984, pp. 91, 93). 13. Bougie. See Pic des Singes National Park. Bouhachem. See Bouhassim, Jebel. Bouhassim, Jebel, 1300 m; Rif, MOROCCO; 35°14'N, 5°26'W reported before 1933 by A. Cabrera (1932, p. 202). Observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 253). Observed 1973-1975 by D. M. Taub (1977, p. 112). Observed Jul-Sep. 1980 by J. E. Fa (1987, "p53; cf.’ Favet al_; 1984, p: 86). Observed Oct._Nov. 2004 by S. S. Waters et al. (2007, p. 107). 33. Bou Iblane, Jebel, between El Mers and Missour; Eastern Moyen Atlas, MOROCCO; 33°39'N, 4°06'’W; reported present before 1932 by L. Joleaud (1933, p. 852). Reported locally ex- tinct before 1978 by D. M. Taub (1977, pulis): 26: Bou Igaouerh, Jebel: MOROCCO; Central Moyen Atlas, 32°57'N, 5°16’W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 94). 13. Bou Jirirh, Central Moyen Atlas, MOROCCO: 33°30'N, 5°09'W; observed Oct. 1980—Oct. 1981 by G. R. Drucker (1984, pp. 136, 138). 25s Bou Tferda. See Abid, Oued el. Boutferda vicinity, Ait Abdi; Southern Moyen Atlas, MOROCCO; ca. 32°22'N, 5°51'W; reported in 1931 by L. Joleaud (1933, p. 852). 9. Bouzega, Djebel. See Bou Zegza, Djebel. Bou Zegza, Djebel, Grande Kabylie, ALGERIA; ca. 36°36’N, 3°27’E; observed in 2005 by M. Oudahmane and B. Asselah (2005, p. 31), locality recolonized by M. sylvanus. 49. Buhasen. See Bouhassim, Jebel. Bu Haxen. See Bouhassim, Jebel. Buseitun. See Buzeitune, Jebel. Buzeitune, Jebel, Beni Hozmar, 1300 m; Rif, MOROCCO; ca. 35°30'N, 5°20’W: observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 253; cf. Fag 1983a,. p. 79). Reported before 1985 by J. E. Fa et al. (1984, pp. 85, 86). Be. Carbon, Cap. See Bouak, Cap. Cedre Gouraud, 1800 m; Central Moyen Atlas, MOROCCO; 33°25'N, 5°10'W; observed Oct. 1994-Sep. 1995 by I. Machairas (Machairas et al., 2003, p. 186). Observed 29 Sep. 2006 by J. Fooden. 23. Ceuta. See Moussa, Jebel. Chabet El Akra. See Akra, Chabet El. Chechaouene (?=Xauen), Jebel; Rif, MOROC- CO; 35°10'N, 5°16’W; observed 1955-1975 by Mr. Valverde, subsequently extinct at this locality (Alvarez & Hiraldo, 1975, p. 258). 34. Cheminot, Djebel Dyjurdjura; Grande Kabylie, ALGERIA; ca. 36°26'N, 4°07'E; blood sam- ples collected before 2000 by F. von Segesser et al (1999: p35) O- Chifa. See Chiffa. Chiffa, Gorges de la; Chiffa; ALGERIA; 36°30’N, 2°45’E; collected before 1940 by unknown collector (Napier, 1981, p. 6); BM(NH), | (not seen; cranial measurements kindly provided by D. Brandon-Jones). 46. ?Chiffa, Gorges de la (locality uncertain); Chiffa, ALGERIA; 36°30’'N, 2°45’E; date and collec- tor unknown (Napier, 1981, p. 6); University Museum of Zoology, Cambridge, 2 (skulls only; not seen). 46. Chiffa, Gorges de la, ca. 15 km S of Blida, 1630 m;_ Chiffa, ALGERIA; ca. 36°21’N, FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 45 2°45'E; observed 28 Oct. 1888 by P. L. Sclater (1888, p. 30). Reported in 1925 by L. Joleaud (1926, p. 129; cf. 1933, p. 852). Observed in 1975 by D. M. Taub (1977, pp. 119, 121; cf. Fa et al., 1984, p. 99). 47. Chrea, 5 km S of Blida, National Park of the Cedars; Chiffa, ALGERIA; 36°25'N, 2°53’E; reported present ca. 1954-1973 and absent in 1974 by J. Chitty and J. P Thomas (Taub, 1984a, p. 75). 48. Collo, Kabylie de, vicinity; region unknown, ALGERIA; ca. 37°00'N, 6°30'E; reported present in 1931 by L. Joleaud (1933, p. 853). Reported to be beyond eastern limit of species distribution in 1978 by D. M. Taub (1978a, p. 249; 1984a, p. 75). 59. Constantina. See Constantine. Constantine; Constantine; ALGERIA; 36°22’N, 6°37’'E; reported before1527 by Leo Africanus (1896 edition, p. 948). Reported to be beyond eastern limit of species distribution in 1978 by D. M. Taub (1978a, p. 249; 1984a, p. 75). 58. Dar El Oued, SSW of Jijel, 800-1200 m; Petite Kabylie, ALGERIA; ca. 36°40'N, 5°35’E; observed 1973-1975 by D. M. Taub (1977, pp. L22e 36. Darguina. See Darguinah. Darguinah; Petite Kabylie, ALGERIA; 36°34'N, 5°18'E; reported by forestry officials to be locally extinct before 1962 (D. M. Taub, 1977, p. 122; cf. Fa et al., 1984, p. 105). 54. Derti-Na, N of, Oued Laou; Rif, MOROCCO; ca. 35°22'N, 5°17'W; observed beforel1976 by F. Alvarez and F. Hiraldo (1975, p. 253; cf. Fa., 1983a, p. 79). 32. Djebel Bouzega. See Bou Zegza, Djebel. Djidjelli. See Ziama Mansouria. Djijel. See Adendoun, Djebel; Dar El Oued; Guerrotch, Forét Domaniale de. Djurdjura, Djebel; Grande Kabylie, ALGERIA; ca. 36°27'N, 4°15’E; tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). SJ. Djurdjura, Djebel. See Ait Oubane, Forét d’; lirilt Aimet; Tala Guilef; Tigounatine; Tikjda. Edough, Djebel, Annaba vicinity; Annaba, AL- GERIA; 36°52'N, 7°39’E; reported present in historic times and locally extinct before 1932 by L. Joleaud (1931b, p. 154). 62. El Hammam, 1200-1450 m; Central Moyen Atlas, MOROCCO; 33°28’N, 5°13’W; ob- served 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, p. 91). 22. 46 El Haouz region; Rif, MOROCCO; ca. 35°30'N, 5°20'W; observed Jan. 2005 by A. El Harrad (Waters et al., 2007, p. 107). 32. El Kelaa; Rif, MOROCCO; ca. 35°10'N, 5°15'W; reported present before 1933 by A. Cabrera (1932, p. 202). Reported locally ex- tinct before 1984 by J. E. Fa (1983a, pp. 69, 79). 34. El Ksiba vicinity, 1800-2000 m; Southern Moyen Atlas, MOROCCO; ca. 32°35'N, 6°02'W; observed 1973-1975 by D. M. Taub (1977, p. 119; cf. Fa et al., 1984, p. 96). 10. Farda, Oued, Rhomara tribal area; Rif, MO- ROCCO; 35°13'N, 4°10'W; frequent local reports in 1981-1982 to P. T. Mehlman (1984, pp. 166, 170). 34. Fath Lemhar, Jebel El Haouz survey area; 700 m; Rif, MOROCCO; ca. 35°39’N, 5°29'W; observed Jul.Sep. 1980 by J. E. Fa (1982, p. 52; cf. Fa et al., 1984, p. 86). 3/. Gargide Chiffa. See Chiffa, Gorges de la. Gibraltar, ca. 400 m; Gibraltar, U.K.; ca. 36°08'N 5°21’W; introduced repeatedly, including in or before 1740 (cf. Busk,1877, p. 130; Kenyon, 1938, pp. 1, 4, 111, 116; Fa, 1984b, p. 266; Modolo et al., 2005, p. 7397). Collected before 1865 by unknown collector; zmuz, 1. Collected before 1910 by unknown collector (cf. Napier, 1981, p. 6); BM(NH), 2 (skins only). Collected 11 Dec. 1930 by unknown collector; UsNM, 1. Collected Feb. 1937 by unknown collector; FMNH, 3. Collected 11 Nov. 1949 by R. Henry; IRSN, | (skeleton only). 29. Gibraltar, N end of mountain; Gibraltar, U.K.; ca. 36°09'N, 5°21'W; unfossilized skulls, ten- tatively identified as M. sy/vanus (introduced), collected before 1798 by miners (Imrie, 1798, pp. 198, 201), museum unknown, 2 (skulls only). 29. Gorges de la Chiffa. See Chiffa, Gorges de la. Goundafa; Haut Atlas, MOROCCO; 31°00'N, 8°05'W; reported present before 1926 by L. Joleaud (1926, p. 129). Reported locally ex- tinct in 1973-1975 by D. M. Taub (1977, hi). 2 Gouraya, Djebel. See Pic des Singes National Park. Gouroud. See Cedre Gouraud. Guerrotich, Forét Domaniale de, SSW of Jijel, 800-1200 m; Petite Kabylie, ALGERIA; 36°42’N, 5°37’E; observed 1973-1975 by D. M. Taub (1977, p. 122; cf. Fa et al., 1984, p. 107). 56. FIELDIANA: ZOOLOGY Guessig, Mount, Djurdjura National Park, 1480— 2100 m; Grande Kabylie, ALGERIA; 36°27'N, 4°08'E; observed Apr. 1983—Jul. 1988 by N. Ménard et al. (1990, pp. 167, 169). 5/. Haounet, Forét dE], vicinity, Ait Abdi; Southern Moyen Atlas, MOROCCO; ca. 32°13'N, 5°56'W; reported in 1931 by L. Joleaud (1933. py 852).9. Haouz, Jebel El. See Fath Lemhar. Herrata; region unknown, MOROCCO; not located; collected 20 Apr. 1904 by W. Schliter; USNM, |. Not mapped. Hotel, Djebel Djurdjura; Grande Kabylie, AL- GERIA; ca. 36°26'N, 4°08’E; blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). SJ. Icetcifene, Djebel, ca. 2 km from Tigounatine, Djebel, Dyebel Dyurdjura, 1600-2100 m; Grande Kabylie, ALGERIA; ca. 36°27'N, 4°08'E; observed 1983-1985 by N. Ménard and D. Vallet (1986, p. 174). 5/. Icetcifere. See Icetciféne, Djebel. Idjberten. See Ijbertene, Arhbalou. Idrous, Jebel; Southern Moyen Atlas, MOROC- CO; ca. 32°08'N, 6°04'W; reported in 1931 by L. Joleaud (1933, p. 852). 8. Ifrane, 1650 m; Central Moyen Atlas, MOROC- CO; 33°32'N, 5°06’W; observed 1973-1975 by DIM. Taub (977s \p: 116)/.23. Ighrem, [Jebel], Kasba Tadla-Beni Mellal region; Southern Moyen Atlas, MOROCCO; ca. 32°20'N, 6°15'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 96). 7. lirilt Aimet, Djebel Dyurdjura, 1750-2300 m; Grande Kabylie, ALGERIA; not precisely located; ca. 36°27’N, 4°15’E; observed 1982-— 1983 by N. Ménard (in Fa et al., 1984, p. 103). Not mapped. Ijbertene, Arhbalou; Southern Moyen Atlas, MOROCCO; 32°10'N, 5°46’W; reported in 1931 by L. Joleaud (1933, p. 852). 9. Illjida, Oued. See Tilljida, Oued. Imi Ourarn, above Lake Miami, 1600-2000 m; Central Moyen Atlas, MOROCCO; ca. 32°54’N, 5°22’W; observed 1973-1975 by D. M:.. Taub 1977; -p: 118;1ef:, Fa ‘et. al., 1984, pp: 91.194) 15: Immouzer-du-Kandar. See Imouzzer du Kandar. Imouzzer du Kandar, vicinity; Central Moyen Atlas, MOROCCO; ca. 33°44’N, 5°01'W; observed 1973-1975 by D. M. Taub (1978a, p. 246). 24. Itzer, 10-15 km E of; Central Moyen Atlas, MOROCCO; ca. 32°53'N, 4°55'W: observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 95). 19. Itzer, W of; Central Moyen Atlas, MOROCCO; ca. 32°53’N, 5°03’W; observed 1973-1975 by D. M. Taub (1977, p. 118} cf. Fa et al., 1984, pe95).L9: Itzere. See Ttzer. Jemmapes. See Azzaba. Jirrir; Jirigh. See Bou Jirirh. Kaiat, Jebel, 1300m; Rif, MOROCCO; 35°18'N, 5°09'W; observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 254). Ob- served Jul—Sep. 1980 by J. E. Fa (1982, p. 52; cf. Fa et al., 1984, p. 86). 34. Katama. See Ketama. Kebir, Oued El, gorge, S of El Milia; E/ Milia, ALGERIA; ca. 36°40'N, 6°15’E; reported in 1931 by L. Joleaud (1933, p. 853), beyond modern eastern limit of distribution (Taub, 1984a, p. 75). 57. Kelti, Jebel, 1300 m; Rif, MOROCCO; 35°22'N, 5°17'W; observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 253). Observed Jul.— Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 86). 32. Kerrata. See Akra, Chabet El; Babor, Djebel. Kerrouchene; Central Moyen Atlas, MOROC- CO; 32°49'N, 5°20’W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 95). 73. Ketama vicinity, Senhaja de Srair region; Rif, MOROCCO; ca. 34°47'N, 4°40'W; reported present before 1932 by L. Joleaud (1933, p. 852). Reported present in mid-1960s and absent in 1973-1975 by J. Chitty (in Taub, 1977, p. 113; 1984a, p. 73). 38. Khenifra, 22 km E, 1600 m; Central Moyen Atlas, MOROCCO; 32°58'N, 5°27'W;; collect- ed 8 Jul. 1970 by R: E. Vaden; usnm; 2 (including | skull only). /3. Khenifra area, Central Moyen Atlas, MOROC- CO; ca. 32°56'N, 5°40’W; blood samples collected Aug. 1977 (Stanyon et al., 1980, p. 149). 12. Kherrata. See Akra, Chabet El; Babor, Djebel. Kherrata, Gorges de; Petite Kabylie, ALGERIA; ca. 36°29'N, 5°17’E; observed 1982-1991 by N. Ménard et al. (1992, p. 156). Blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). 54. Komuch, El Ksiba vicinity; Southern Moyen Atlas, MOROCCO; ca. 32°35'N, 6°02'W; observed 1973-1975 by D. M. Taub (1977, p. 119: Fa et al.,s19845'p..97). 10: Koucer. See Kousser, Massif du. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 47 Kousser, Massif du; Southern Moyen Atlas, MOROCCO; 32°03'N, 6 02'W; reported in 1931 by L. Joleaud (1933, p. 852). &. Ktama. See Ketama. Lakraa, Jebel, 1600-2160 m; Rif, MOROCCO; 35°08'N, 5°09'W; observed 17 Oct.-27 Nov. 1971 by A. Whiten and T. J. Rumsey (1974, p. 421). Observed beforel976 by F. Alvarez and F. Hiraldo (1975, p. 257). Observed Jul.- Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 86). 34. Lau. See Derti-Na, Sidi Siah. Maggo. See Magoo, Jebel. Magoo, Jebel; Rif, MOROCCO; ca. 35°10'N, 5°15'W; reported present before 1933 by A. Cabrera (1932, p. 202). Reported locally ex- tinct before 1984 by J. E. Fa (1983a, p. 69). 34. Mansouria. See Ziama Mansouria. Mauretania; region unknown, MOROCCO and/ or ALGERIA; ca. 34-37°N, 3°W-5S°E; re- ported before 1527 by Leo Africanus (1896 edition, p. 948). Not mapped. Mauritania. See Mauretania. Merhraoua. See Tahafourt. Miami, Lake. See Arhbalou N’Irkraouen; Imi Ourarn. Michlifene, 2000 m; Central Moyen Atlas, MO- ROCCO; 33°25'N, 5°07'W; observed 1973- 1975 by D. M. Taub (1977, p. 116). 23. Mimejab. See Mimejad. Mimejad; Central Moyen Atlas, MOROCCO; 33°00'N, 5°16’W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 94). /3. Mischliffin. See Michlifene. MOROCCO (locality and region unknown); collected 7 Aug. 1931 by Dr. Lionville; MNHN, 1. Not mapped. Mouga. See Moussa, Jebel. Moussa, Jebel, Anjra massif, W of Ceuta, 800 m; Rif, MOROCCO; ca. 35°54’N, 5°24’W; re- ported before 1932 by L. Joleaud (1933, p. 852). Observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 254). Observed Jul.— Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 86). Observed Oct.—Nov. 2004 by S. S. Waters et al. (2007, p. 107). 30. Muga. See Moussa, Jebel. Musa. See Moussa, Jebel. Nador vicinity; Rif, MOROCCO; ca. 35°00'N, 3°00'W; population introduced in 1985 (Au- lagnier & Thevenot, 1986, p. 56). 43. National Park of the Cedars. See Chrea. North Saqqara; EGYPT; ca 29°53’N, 31°15’E; mummified captives (introduced), ca. 400— 48 0 B.C.E. (Goudsmit & Brandon-Jones, 1999, p. 51). Not mapped. Ouaouezarht. See Ouaouizarht. Ouaouizarht, MF de; Southern Moyen Atlas, MOROCCO; = 32°10'N, 621'W; observed 1973-1975 by D. M. Taub (1977, p. 119; ef. Fa et al., 1984, p. 96). 6. Oued el Rbia. See Oum Er Rbia, Oued. Ouiouane; Central Moyen Atlas, MOROCCO; 33°07'N, 5°22'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 93). 22. Oum Er Rbia, Oued, source; Central Moyen Atlas, MOROCCO; 33°04'N, 5°25'W; ob- served 1973-1975 by D. M. Taub (Fa et al., 1984, p. 93). 22. Ourika Valley. See Ourika, Oued; Tourchte; Zaclia Sti Fatma. Ourika, Jebel; Haut Atlas, MOROCCO; ca. 31°15'N, 7°47'W; reported before 1926 by L. Joleaud (1926, p. 129). 2. Ourika, Oued; Haut Atlas, MOROCCO; ca. 31°15'N, 7°38'W; reported in 1926 by P. Bédé (1927, p. 28). Observed in 1981 by G. R. Drucker (Fa et al., 1984, pp. 80, 99). Tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). 2. Ouzoud, Cascades d’, Oud-el-Abid gorge, Azilal district; Southern Moyen Atlas, MOROCCO; ca. 32°02'N, 6°47'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 97). Observed in 1981 by G. R. Drucker (Fa et al., 1984, p. 97). Reported before 1985 by G. de Turckheim (Taub, 1984a, p. 73; Fa et al., 1984, p. 97). Tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393; cf. Fa et al., 1984, p. 97). 3. Palestro, Gorges de, Bou Zegza; Grande Kabylie, ALGERIA; 36°36’N, 3°35’E; reported in 1925 and 1931 by L. Joleaud (1926, p. 129; 1933, p. 853). Reported absent before 1985 by D. M. Taub (1984a, p. 75). 50. Philippeville. See Stora vicinity. Pic des Singes National Park, Djebel Gouraya, near Bejaia, 600m; Grande Kabylie, AL- GERIA; 36°46’N, 5°05’E; reported before 1527 by Leo Africanus (1896 edition, p. 948). Reported in 1925 by L. Joleaud (1926, p. 129). Observed in 1931 by L. Joleaud (1933, p. 853). Observed in 1975 by D. M. Taub (1977, p. 120; cf. Fa et al., 1984, p. 102). Blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). Tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). 53. FIELDIANA: ZOOLOGY Pompeii; ITALY; 40°31’N, 14°29’E; skeletal fragments of introduced captive, radiocarbon dated 50 B.C.E.-140 C.E. (Bailey et al., 1999, p. 1413). Not mapped. Ras El Ma, 1800 m; Central Moyen Atlas, MOROCCO; 33°28'N, 5°07'W; observed 1973-1975 by D. M. Taub (1977, p. 116). 23. Ras Timedouine, Djebel Dyjurdjura; Grande Kabylie, ALGERIA; ca. 36°27'N, 4°14'E; observed 1982-1983 by N. Ménard (in Fa et al., 1984, p. 103). 5/. Ras Timedouire. See Ras Timedouine. Rass El Ma. See Ras El Ma. Rhiata mountains, SW of Taza; Eastern Moyen Atlas, MOROCCO; ca. 34°10'N, 4°07'W; reported present before 1932 by L. Joleaud (1933, p. 852). Reported locally extinct 1973- 1975 by D. M. Taub (1977, p. 115; 1984a, p73): 28. Riata. See Rhiata mountains. Rif (imprecise locality); Rif, MOROCCO; ca. 35°N, 4°W: tissue samples collected before 2006 by L. Modolo et al. (2005, p. 7393). Not mapped. Rond-Point des Cédres; Theniet el Had, AL- GERIA; 35°52’N, 1°57'E; reported present in 1954 and absent in 1974 (J. Chitty and J. P Thomas in Taub, 1984a, p. 75). 44. Ruisseau des Singes. See Chiffa, Gorges de la. Seheb, ca. 15 km SE of Azrou, 1700 m; Central Moyen Atlas, MOROCCO; ca. 33°20'N, 5°05'W; observed 1973-1975 by D. M. Taub (1977, p. 116; cf. Favet al., 1984, p..91)..2/. Senoual region, 1800 m, Central Moyen Atlas, MOROCCO; ca. 32°58'N, 5°13’W; observed 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, p. 91). /3. Setti Fatma. See Zaclia Sti Fatma. Sidichmim, Beni Hozmar; Rif, MOROCCO; ca. 35°30'N, 5°20'W; observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 253; cf. Fa., 1983a, p. 79). 32. Sidi Mguid vicinity; Central Moyen Atlas, MOROCCO; ca. 33°13’N, 5°16’W; reported before 1932 by L. Joleaud (1933, p. 852). Observed 1973-1975 by D. M. Taub (1977, p. 116; cf. Fa et al., 1984, p. 90). 22. Sidi M’Guild. See Sidi Mguid. Sidi Salah, Jebel, 1300 m; Rif, MOROCCO; 35°17'N, 5°09'W; observed Jul.—Sep. 1980 by J. E. Fa (1982) p. 59;:cf. Favet al:, 1984, p: 86). 34. Sidi Siah, at confluence of Oued Laou and Uad Talambot, Rif, MOROCCO; 35°17'N, 5°14'W; observed before 1976 by F. Alvarez and F. Hiraldo (1975, p. 253). 34. Sidi Yahia ou Youssef. See Sidi Yahya ou Youssef. Sidi Yahya ou Youssef, 8-10 km W of, 1900 m; Southern Moyen Atlas, MOROCCO; ca. 32°24'N, 5°28’W; observed 1973-1975 by D. Mi: Taub: (1977, p: 119;cf Fa ‘et al, 1984 pes6)71S: Sidi Yahya ou Youssef, S and W of, 1900 m:; Southern Moyen Atlas, MOROCCO; ca. 32°24'N, 5°22'W; observed 1973-1975 by D. M» faub- (1977, p: 119; cf. Fa et°als 1984, poo}. 1S Sokmane. See Ait Sokhmane forest. Source des Singes, Djebel Djurdjura; Grande Kabylie, ALGERIA; ca. 36°25’N, 4°13’E; blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). SJ. Spain, [southern]; Andalusia, SPAIN: ca. 37°N, 5°W; spurious reports (I Geoffroy, 1851, p. 31; Lydekker, 1893-1894, p. 118; cf. 1916, p. 7; Forbes, 1894, p. 5; cf. Drinkwater in Kenyon, 1938, p. 112). Not mapped. Stora, “Oran”; probably Skikda, ALGERIA; probably ca. 36°54'N, 6°52’E; collector and date unknown (Napier, 1981, p. 6); BMcNH), | (skin only, skull inside). 60. Stora vicinity, W of Skikda; Skikda, ALGERIA; ca. 36°54'N, 6°52’E; reported present in 1925 by L. Joleaud (1926, p. 129). Population reported near extinction in 1931 by L. Joleaud (1931b, p. 154; 1933, p. 853). Population reported extinct before 1978 by D. M. Taub. (197 7 -spsl23). 60: Tabarca. See Tabargqah. Tabarqah; Beja, TUNISIA; 36°57'N, 8°45’E; reported present in historic times by L. Joleaud (1931b, p. 154). 63. Tafechna, ca. 10 km SW of Aguelmane Azigza; Central Moyen Atlas, MOROCCO; ca. 32°54'N, 5°30'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 94). 73. Taffert, at Tine M’lilt, 1700-2000 m; Eastern Moyen Atlas, MOROCCO; ca. 33°45'N, 4°17'W; observed 1973-1975 by D. M. Taub (1977, pedi blts)-25: Tagoulelt vicinity; Southern Moyen Atlas, MO- ROCCO; ca. 32°32'N, 4°48’W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 96). J8. Taguelft region. See Asker; Azrar; Tsemair. Tahafourt, 4 km S of Merhraoua, 1400-1700 m; Eastern Moyen Atlas, MOROCCO; 33°52'N, FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 49 4°04'W: observed 1973-1975 by D. M. Taub (1977, pp. 111, 115; cf. Fa et al., 1984, p. 80). Tahar Souk, N of; Rif, MOROCCO; ca. 34°40'N, 4°15'W; reported before 1958 to M. Ricard by Moroccan informants (Panouse, 1957, p. 28). 25. Takoucht, Djebel; Petite Kabylie. ALGERIA; 36°31'N, 5°13’E; population reported extinct before 1962 by forestry officials (Taub, 1977, p. 122; cf. Fa et al., 1984, p. 105). 54. Talaat Ardhousse, 1300 m; Rif, MOROCCO; ca. 35°16'N, 5°08'W; observed Jul.-Sep. 1980 by J. E. Fa (1982, p. 59). 34. Tala Guilef. See Tala Gulief. Tala Gulief, Djebel Djurdjura, 1750-2300 m; Grande Kabylie, ALGERIA; ca. 36°29'N, 4°00'E; observed 1982-1983 by N. Ménard (in Fa et al., 1984, p. 103). Blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). 5/. Tala Kitane, Akfadou vicinity, 900-1278 m; Grande Kabylie, ALGERIA; 36°41'N, 3°34’E; observed 1982-1983 by N. Ménard (in Fa et al., 1984, p. 104). 50. Talambot. See Sidi Siah. Tala Rana, Djebel Djurdjura; Grande Kabylie, ALGERIA; ca. 36°25'N, 4°15’E; blood sam- ples collected before 2000 by F. von Segesser et al. (1999, p. 435). 5/. Talassemtane, Rhomara tribal area, 2100 m; Rif, MOROCCO; ca. 35°09'N, 5°07'W; observed Jul.Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 86). Observed 1981-1982 by P. T. Mehlman (1984, p. 170). Observed Oct.— Nov. 2004 by S. S. Waters et al. (2007, p. 107). 34. Tamjilt, below, 1700-2000 m; Eastern Moyen Atlas, MOROCCO; 33°39'N, 4°00’E; ob- served 1973-1975 by D. M. Taub (1977, pp. 111, 115). 26. Taourirt Irhil, Forét de, 500-1600 m, Grande Kabylie, ALGERIA; 36°41’N, 4°40’E; ob- served 1973-1975 by D. M. Taub (1977, p. 122; cf. Fa et al., 1984, p. 104). 52. Targa vicinity, Beni Ziat area; Rif, MOROCCO; ca. 35°24'N, 5°01'W; reported before 1932 by L. Joleaud (1933, p. 852; cf. Fa, 1983a, p. 79). 36. Tarhzirt. See Ouzoud, Cascades d’. Tasaot, Jebel, Rhomara tribal area, 1700- 2000 m; Rif, MOROCCO; 35°14'N, 5°06'W; observed 1973-1975 by D. M. Taub (1977, p. 111). Observed before 1976 by F. Alvarez 50 and F. Hiraldo (1975, p. 254). Observed Jul.— Sep. 1980 by J. E. Fa (1982, p. 59; cf, Fa et al., 1984, p. 87; Mehlman, 1984, p. 170). 34. Tazaot. See Tasaot, Jebel. Tazaoute. See Tasaot, Jebel. Tazonte, Djebel. See Tasaot, Jebel. Tétouan vicinity; Rif, MOROCCO; ca. 35°34'N, 5°22'W; reported present before 1926 by L. Joleaud (1926, p. 129). Reported locally ex- tinct before 1983 by J. E. Fa (1982, p. 48). 32. Theniet el Had. See Rond-Point des Cedres. Tidiquin. See Tidirhine. Tidirhine, Jebel; Rif, MOROCCO; 34°S1’N, 4°31'W; reported present in mid-1960s and locally extinct subsequently by J. Chitty and local forestry official (Taub, 1984a, p. 73). 39. Tighret forestry post, 25 km NW of Miliana; Chiffa, ALGERIA; ca. 36°25’N, 2°00’E; re- ported present in 1954-1973 and locally extinct in 1974 by J. Chitty and J. P. Thomas (Taub, 1978a, p. 249; 1984a, p. 75). 45. Tigounatine, Djebel Dyjurdjura, 1750-2300 m; Grande Kabylie, ALGERIA; 36°27'N, 4°08'E; observed 1973-1975 by D. M. Taub (1977, p. 120; cf. Fa et al., 1984, p. 102). Observed 1983-1984 by N. Ménard et al. (1985, p. 66). Blood samples collected before 2000 by F. von Segesser et al. (1999, p. 435). SJ. Tiguerouguine, Plateau d’Ajdir; Central Moyen Atlas, MOROCCO; ca. 32°55'N, “S°2595 observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 94). /3. Tijjida, Oued. See Tilljida, Oued. Tikjda, Djebel Djurdjura, 1200-1900 m; Grande Kabylie, ALGERIA; 36°27'N, 4°08’E; ob- served 1983-1985 by N. Ménard et al. (Fa et al., 1984, p. 103; Ménard et al., 1986, p. 35). St. Tilljeda, Oued. See Tilljida, Oued. Tilljida, Oued, Rhomara tribal area, 400 m; Rif, MOROCCO; 35°11'N, 5°06’W; observed Jul.— Sep. 1980 by J. E. Fa (1982, p. 58; cf, Fa et al., 1984, p. 87; Mehlman, 1984, p. 170). 34. Tillouguit N’Ait Isha, S of; Southern Moyen Atlas, MOROCCO; ca. 32°02’N, 6°13’W; observed 1973-1975 by D. M. Taub (1977, p. 119; cf. Fa et al., 1984, p. 97). 5. Tilougguite. See Ouzoud, Cascades d’; Tillouguit N’Ait Isha. Tine M’lilt. See Taffert. Tirizene, Djebel. See Tisirene, Jebel. Tisirene, Jebel, above Boureit, 1800 m; Rif, MOROCCO; 35°01’N, 4°55’W; observed 1973-1975 by D. M. Taub (1977, pp. 111, FIELDIANA: ZOOLOGY 112). Reported in 1984 by J. E. Fa et al. (1984, p. si: cl. Fa, 19836, p: 62). 35. Tisouka, Jebel, Rhomara tribal area, 2100 m; Rif, MOROCCO; 35°10'N, 5°13’W; observed Jul.Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 87; Mehlman, 1984, p. 170). 34. Tissouka, Djebel. See Tisouka, Jebel. Tiz Ifmi. See Tizi Ifri. Tizi Ifri (?=Tiz Ifmi); Rif, MOROCCO; 34°51'N, 4°15'W; reported present in mid-1960s and locally extinct subsequently by J. Chitty and local forestry official (Taub, 1984a, p. 73). 40. Tizi M’Isly, N_ of; Southern Moyen Atlas, MOROCCO; ca. 32°28'N, 5°46’W; observed 1973-1975 by D. M. Taub (1977, p. 119). 74. Tizirane, Djebel. See Tisirene, Jebel. Tounfit, 15-20 km SSE of; Southern Moyen Atlas, MOROCCO; ca. 32°20’N, 5°10’'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p. 96). /6. Tounfit, N and W of; Southern Moyen Atlas, MOROCCO; ca. 32°30'N, 5°15'W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, p96): 17. Tounfite. See Tounfit. Tourchte, Oued Ourika; Haut Atlas, MOROC- CO; 31°15'N, 7°38’W; multiple reports in Jul.— Aug. 1968 received by J. M. Deag and J. H. Crook (1971, p. 186). 2. Tsemair, Taguelft region; Southern Moyen Atlas, MOROCCO; ca. 32°15'N, 6°08’W; observed 75-19 75\'by) D. M. Taub: (1977, pr 119). Possibly locally extinct 1974-1982 (Fa et al., 1984, p. 96). 8. Xauen. See Chechaouene. Zaclia Sti Fatma, 50 km SSE of Marrakesh, ca. 1900 m; Haut Atlas, MOROCCO; 31°18'N, 7°47'W; collected 15 Aug. 1968 by unknown collector . (Napier, 1981, p. 6; cf..Deag: & Crook, 1971, p. 186); BMcNH), 1 (skull only; not seen). 2. Zaclia Sti Fatma, S of, Oued Ourika, 1900 m; Haut Atlas, MOROCCO: ca.” 31° 13’N; 7°42'W; observed Jul—Aug. 1968 by J. M. Deag and J. H. Crook (1971... pr1s6A ck Napier, 1981, p. 6). 2. Zad region, 2300 m; Central Moyen Atlas, MOROCCO; ca. 33°01'N, 5°04’W; observed 1973-1975 by D. M. Taub (Fa et al., 1984, Pp 95)r20; Zaghouan. See Zaghwan. Zaghwan; Zaghwan, TUNISIA; 36°24’N, 10°09'E; reported present in historic times (Joleaud, 1931b, p. 154). 64. Zaouia de Ifrane; Central Moyen Atlas, MO- ROCCO; ca. 33°14'N, 5°25'W; observed 1973- 1975 by D. M. Taub (Fa et al., 1984, p. 94). 22. Zarka cascade vicinity, Beni Hozmar area; Rif, MOROCCO; ca. 35°28'N, 5°20'W; reported before: 1932 by LL’ Joleaud (1933, pr852)i a2: Ziama Mansouria, E of, Jijel, W of; Petite Kabylie, ALGERIA; ca. 36°40'N, 5°30’E; reported in 1931 by L.- Joleaud (1933, p. 853). 56. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE Sil Appendix III. Centrum Length and Vertebral Count of Caudal Vertebrae in Macaca sylvanus Length (mm) of indicated caudal vertebrae liferred Canial vente Specimen No. 1 2 3 4 5 count! Juvenile females FMNH 47410 8.0 i ils m- = < 3 IRSN 9165Q 10.9 15.2° — a IRSN 9071 9.6 7.8 7.1 a — 3 zmuz 11602* 8.1 8.1 11.9 1.4 = 4 zsBs 1959/4 10.3 fe id a Ma a Juvenile males IRSN 91650" 10.5 75 x os 3 IRSN 16701 10.5 10.9 10.8 — = 3 IRSN 16696 7.8 8.1 8.0 a =e 3 ns 1552 9.7 10.3 1) us — 3 ZMB 8264 8.5 10.5 9.2 ha! os 3 Adult females FMNH 47409 Bi >5.9° = = ea ~) IRSN 9295 10.9 10.4 5.3 m~ aa >4 IRSN 17073 11.1 8.5 7.2 — eee 3 IRSN 17058 9.9 8.7 6.3 — = 3 IRSN 16871 10.8 11.7 14.2° a >4 RMNH 9c 11.9 13.1 11.4 13.1 3.0 5 USNM 476786 10.0 LE 4.0 =e = 2 zmB 16492 12.0 18.9° ws a8 33 Means° 10.96 10.68 6.84 13.1 3.0 — Adult males FMNH 47398 16.2 v4 Wel vil ats >3 mcz 7098 13.1 Bits: 10.3 — — 3 RMNH 7a 13.2 18.9° =e a! 3 USNM 476785 ee 8.9 m? map es >3 ZMB 8271 (ies! 11.9 6.1 ae — 3 Means° 13.30 10.70 8.20 = — — ' Supplementary counts from literature (age, sex, and/or measurements of specimens usually not specified): Vrolik, 1841, p. 10—2 caudal vertebrae; Mivart, 1865, p. 583—3, 4, and 4 caudal vertebrae; Giebel and Leche, 1874-1900, p. 241—4 caudal vertebrae; Didier and Rode, 1936, p. 3—generally 2 caudal vertebrae; A. W. Todd in Schultz and Straus, 1945, p. 613—2 caudal vertebrae. > Terminal vertebra/vertebrae missing. * Cd2 and Cd3 fused; values excluded from calculation of means. * Sex uncertain. ° Vertebra broken. ® For adult specimens only. 52 FIELDIANA: ZOOLOGY Appendix IV. Group Size and Adult Sex Composition in Macaca sylvanus Populations Group size Adult composition No. of Mini- — Maxi- MIF _ Refer- Sample area groups Mean mum mum Males Fe-males _ ratio ences Morocco: Moyen Atlas Afennourir 9 37.8 — — == — 1.21 1 Ain Kahla 1 39 a — 4) 9 ; 2 Ain Kahla 16 24.1 12 39 66 94 0.70 3 Ain Kahla 1 35 — — — = — 4 Bou Jirirh 4 1 40 ao — — — — 5 Ceédre Gouraud~ 3 23 — — — — 0.96 1 Oued-el-Abid 1 18 — 3 5 0.60 6 Sidi Mguid l 15 _ — — — — 7 Regional summary 33 28.6 12 39 76 108 0.70° o— Morocco: Rif Anjera: Moussa, Jebel* ] f2 — — l 2 0.50 8 Beni Hessane: Bouhassim* 5 13.6 8 27 15 1] 1.36 8 Beni Zaid: Kelti, Jebel 1 12 — — 3 2 1.50 8 Beni Zaid: Sidi Salah, Jebel 1 14 — — p 3 0.67 8 Beni Zaid: Talaat Ardhousse 1 10 — — 2, 4 0.50 8 Rhomara: Adelma, Oued 2 itil 10 12 6 6 1.00 8 Rhomara: Lakraa, Jebel 4 16.8 12 28 1] 11 1.00 8 Rhomara: Lakraa, Jebel 2 Dies DS 30 18 9 2.00 9 Rhomara: Talassemtane* 1 27 — = 4 4 1.00 8 Rhomara: Talassemtane 6 27.0 i2 59 29° 40 0.72 10 Rhomara: Tazoute, Jebel 1 19 = — 4 4 1.00 8 Rhomara: Tisouka, Jebel 2) 13)5) 7 20 5 5 1.00 8 Regional summary 27 18.3 ip 59 100 101 0.99 — Algeria: Grande Kabylie Akfadou® l 53 = = = 1.00 11 Djurdjura: rocky mountains’ 4 45.2 36 58 -— — 0.89 12 Djurdjura: Tigounatine® 1 88 — -- — 1.00 11 Djurdjura: Tikjda’ D 40.0 36 44 ~14 ~17 ~0.82 13 Regional summary 8 50.2 36 88 ~14 ~l7 ~~ ~0182 _ Composite 68 pA ak 7 88 90 ~226 ~0.84 — ' Key to references: 7. Machairas et al., 2003, p. 189; Camperio Ciani and Machairas, 2003, p. 125. 2. Taub, 1980b, p. 290. 3. Deag, 1984, p. 119. 4. Ménard and Qarro, 1999, p. 124. 5. Drucker, 1984, p. 139. 6. Fa, 1986b, p. 32. 7. Hammerschmidt et al., 1994, p. 279. 8. Fa, 1982, p. 59. 9. Whiten and Rumsey, 1974, p. 423. 10. MehIman, 1989, p. 272. /7. Ménard and Vallet, 1993b, pp. 105, 108. 72. Ménard et al., 1990, p. 169. 73. Ménard et al.) 1986; p. 37. * Alternate locality name: Gouroud. > Excludes undocumented Afennourir and Cédre Gouraud ratios. + New censuses of these groups were conducted in 2004 (Waters et al., 2007, p. 107). ° Focal group apparently includes 11 (not 12) adult males. © 1990 census; in 1983 group size was 33. ’ Group size in this area is unstable—subject to subgroup fusion and fission (cf. Ménard et al., 1985, p. 72; Menard, 2002, pp. 96, 98). * 1988 census; in 1983 group size was 38, and in 1989 the group subdivided into three daughter groups. ? December 1985 census; excludes data for group Tl (=Tigounatine group). FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 53 Appendix V: Coordinates, G M eological Age, and Reported Identification of Circum- Mediterranean Macaque Fossils Coordinates Fossil localities (approximate) Epoch Remarks” References® ALBANIA Gajtan 42°03'N 19°34’E Pleistocene Cf. M. sylvanus pliocena; Mid-Pleist. 1 ALGERIA Afalou bou Rhummel 36°30'N 5°20'E Pleistocene MM. cf. sy/vanus; Late Pleist. a Menacer (Marceau) 36°30'N 5°00'E Miocene ? Macaca sp.; probably Late Mio., ca. 7 Ma 25:3} GAS Tamar Hat 36°30'N 5°20'E Pleistocene MM. cf. sy/vanus; Late Pleist. ya. Taya, Djebel, grottos* 36°30'N 7°08'E Pleistocene M. sylvanus 6 Traras, Monts des, 35°02'N 1°40'W Pleistocene M. sylvanus; Mid- or Nedroma Late Pleist., 0.2 Ma 4, 5/6, 78 AUSTRIA Deutsch-Altenburg” 48°10'N 16°55'E Pleistocene MM. sy/vanus; Early Pleist. 9 Kugelsteinhohle, 47°13'N 15°20’E Pleistocene MM. sy/vanus group; Late Pleist., 22 km N of Graz probably Eemian 10 CROATIA Sandalja 45°0S'N 14°00'E Pliocene M. cf. sylvanus; Mid-Villafranchian 2:3; 4088 CZECH REPUBLIC Zlaty Kun, Cave C718 49°35'N 14°45'E Pleistocene MM. sylvanus; Mid-Pleist. 2, 3, 4 EGYPT Wadi Natrun 30°25’N 30°13’E Miocene M. libyca; Late Mio., ca. 6 Ma 2; 3,48 ENGLAND, U.K. Grays Thurrock 51°29’N 0°20’E Pleistocene Macaca sp.; probably late Middle Pleist. 2, 3:83, 1] Hoxne 52°21 N 1°12"E Pleistocene Macaca sp.; Mid-Pleist. 12 Swanscombe (Barnfield 51°26'N 0°18’E Pleistocene Macaca sp.; Mid-Pleist. Pit) 1] West Runton 52°45'N 1°05’E Pleistocene = Macaca sp.; Mid-Pleist. 2, S54 aa FRANCE Balaruc-2 43°35'N 3°45’E Pliocene M. cf. sylvanus; Early Villafranchian 2, 4, 11 Caune de l’Arago 42°33'N 3°00’E Pleistocene MM. sy/vanus, Mid-Pleist, 0.23 Ma 13 Montpellier 43°38'’N 3°53’E Pliocene M. cf. sylvanus; Early/Mid-Ruscinian 2, os eee 1] Montsaunés 43°20'N 0°55’E Pleistocene M. cf. sylvanus; Mid-Pleist. 2, Aye Orgnac-3 43°50'N 5°00’E Pleistocene Macaca sp.; Mid-Pleist. 3, 10 Saint-Esteve-Janson 43°30'N 5°40’E Pleistocene Macaca sp.; Mid-Pleist. 2, 3, Ste Saint-Vallier 45°10'N 4°50’E Pliocene M. cf. sylvanus; Mid-Villafranchian 2, 3 4am Senéze 44°30'N 3°00’E Pleistocene UM. cf. sy/vanus; Late Villafranchian 2, 3,'4,ae 54 FIELDIANA: ZOOLOGY Fossil localities Vallonet GEORGIA Kudaro-1 GERMANY Gundersheim Heppenloch Hohensiilzen Hunas Mosbach-2 Untermassfeld, 2 km S of Meiningen Voigtstedt GREECE Tourkobounja (Athens) HUNGARY Beremend-4 Csarnota-2 Somssichhegy 2 ISRAEL Ubeidiya ITALY Bristie II (Carso Triestino) Cavo Pompi Figari, Capo (Sardinia) Fontana Ranuccio Fornace RDB, Villafranca d’ Asti Grotta degli Orsi Volanti Monte Peglia Monte Sacro (=‘‘Rome’’) Mugello Orciano Pietrafitta Pofi Appendix V: Continued. Coordinates (approximate) 43°45'N 7°40’E 42°30'N 43°40'E 49°35'N 10°45’E 48°1S'N 9°25'E 50°00'N 8°15’E 49°30'N 11°32’E 49°21'N 9°09'E 50°33’N 10°25'E 50°40'N 11°40'E 38°00'N 23°40’E 45°45'N 18°25’E 45°45'N 18°25’E 45°52'N 18°26'E 32°05'’N 36°05’E 45°40'N 13°45'E 41°40'N 13°20'E 41°00'N 9°35’E 41°45'N 13°10'E 44°54'N 8°12’E 42°14'N 14°09’E 42°45'N 12°10’E 41°56'N 12°32’E 43°50'N 10°5S5’E 43°28'N 10°30’E 42°55'N 12°20’E 41°34'N 13°25'E Epoch Pleistocene Pleistocene Pliocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pliocene Pliocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pliocene Pleistocene Pleistocene Pleistocene Pliocene Pliocene Pleistocene Pleistocene Remarks” Macaca sp.; Mid-Pleist. Macaca sp.°; interglacial, ca. 0.350 Ma M. cf. sylvanus; Early Villafranchian M. cf. sylvanus; Mid-Pleist. M. cf. sylvanus; Mid-Pleist. cf. M. sylvanus pliocena; Mid- Pleist. M. cf. sylvanus; Mid-Pleist. M. sylvanus; late Early Pleist., 1.07 Ma M. cf. sylvanus; Mid-Pleist. Macaca sp.; Mid-Pleist. M. sylvanus; Early Villafranchian M. sylvanus; Late Ruscinian cf. M. sylvanus pliocena; Mid- Pleist. M. sylvanus; Mid-Pleist. M. cf. sylvanus; Mid.-Pleist. M. cf. sylvanus; Mid.-Pleist. M. majori; Early to Mid-Pleist. M. cf. sylvanus; Mid-Pleist., 0.458 Ma M. cf. sylvanus; Early Villafranchian M. cf. sylvanus; Late Pleist., probably Eemian M. cf. sylvanus; Mid-Pleist., Cromerian G M. cf. sylvanus; Early Pleist.? M. cf. sylvanus; Mid- Villafranchian M. cf. sylvanus; Late Plio. M. cf. sylvanus; Early Pleist. cf. M. sylvanus pliocena; Mid- Pleist. FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE References* yee er Pa | pes ie As) 2,3, ae ya Page cs 55 Appendix V: Continued. Fossil localities San Vito di Leguzzano Torre in Pietra Tuttavista, Monte Val d’Arno, Upper Valdemino (Borgio Verezzi) Zoppega II, Soave LIBYA Sahabi area MOROCCO Chrafate Ez Zarka NETHERLANDS Steyl’ Tegelen’ ROMANIA Betfia-2 [also infillings 9 and 13] SLOVAKIA Gombasek Vcelare 2 SPAIN Almenara-M (=Casablanca-M) Ambrona Cova Bonica Cova Negra La Puebla de Valverde Solano del Zamborino TUNISIA Ain Brimba Ichkeul, Garaet (lake) ' See Fig. 9. Coordinates (approximate) 45°45'N 11°40'E 41°55'N 12°30'E 40°22'N 9°41'E 43°45'N I1°10°E 44°10'N 8°20’E 45°45'N 11°45’E 30°00'N 21°00'E 33°50'N 5°05'W 34°25'N 5°45'W 51°20'N 6°08’E 51°19’N 6°09"E 46°05'N 22°50’'E 48°50'N 21°35’E 48°35'N 20°48’E 39°44'N 0°13'W 40°50'N 2°25'W 41°25’N 2°15'E 38°53'N 0°08'W 40°13'N 0°56'W 36°50'N 2°50'W 36°45'N 8°30’E 33°40'N 8°50’E Epoch Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Miocene? Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Miocene Pleistocene Pliocene Pleistocene Pliocene Pleistocene Pliocene Pliocene Remarks” M. cf. sylvanus; Mid-Pleist. M. cf. sylvanus; Late Pleist., probably Eemian M. cf. majori; probably Early Pleistocene M. cf. sylvanus; Late Villafranchian M. sylvanus, Mid-Pleist. M. cf. sylvanus; Mid-Pleist., Cromerian G cf. Macaca; Mio.—Plio. M. sylvanus; Late Pleist. M. sylvanus; Late Pleist. M. cf. sylvanus; age uncertain M. cf. sylvanus; Late Villafranchian M. sylvanus; Mid-Pleist. M. sylvanus; Mid-Pleist. cf. M. sylvanus pliocena; Mid- Pleist. M. cf. sylvanus; Late Mio. Macaca sp.; Mid-Pleist. M. cf. sylvanus; Early Villafranchian Macaca sp.; Late Pleist, beginning last glaciation M. cf. sylvanus; Mid- Villafranchian Macaca sp.; Late Pleist, probably Eemian M. sylvanus; Mid-Villafranchian, 3 Ma M. sylvanus; Ruscinian, 3 Ma References® 2, 5 te * Fossil identifications indicated in this column are those specified in the cited references; Ma = million years ago. * Key to references: /. E. Delson, pers. comm., 11 Jan. 2007. 2. Delson (1974, p. 133). 3. Szalay and Delson (1979, pp. 355, 356). 4. Delson (1973, p. 140). 5. Delson (1980a, pp. 16-19). 6. Joleaud (1926, p. 129). 7. Pomel (1892, p. 157; 1896, p. 11). 8. Geraads (1987, pp. 22, 25). 9. Fladerer (1987, p. 2). 10. Fladerer (1989, p. 25; 1991, 56 FIELDIANA: ZOOLOGY Appendix V: Continued. p. 272). 11. Ardito and Mottura (1987, pp. 34-40). /2. Singer et al. (1982, p. 144). 73. Moigne et al. (2006, p. 793). 14. Maschenko and Baryshnikov (2002, p. 411). 7/5. Kahlke and Gaudzinski (2005, pp. 1208, 1212, 1213). 16. Rook and O’Higgins (2005, p. 169). 77. Rook et al. (2001, p. 188). 78. Mazza et al. (2005, p. 212). 79. Gentili et al. (1998, p. 683). 20. Rook et al. (2003, p. 18). 27. de Heinzelin and El-Arnauti (1987, p. 17); Meikle (1987, p. 126). 22. Ouahbi et al. (2001, p. 641). 23. Kohler et al. (2000, p. 450). 24. Eronen and Rook (2004, p. 327). 25. Delson (1980b, p. 126). * Hammam Meskoutine vicinity. > Ca. 50 km SE of Vienna. ® Delson (1973, p. 140) and Szalay and Delson (1979, p. 356) regarded these fossils as closely related to M. sylvanus; Maschenko and Baryshnikov (2002, p. 411) regarded them as more closely related to Asian macaques. 7 Possibly two names for the same locality. : Probably <0.100 Ma (E. Delson, pers. commun., | Jan. 2007). FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE ey Note Added in Proof For further discussion relevant to dessication of the Mediterranean Sea (see above, Evolution and Dispersal), consult the following: Roucny, J, M., J.-P. Suc, J. FerRANDINI, AND M. FERRANDINI 2006. Editorial: The Messinian Salinity Crisis revisited. Sedimentary Geology, 188-189: 1-8. Suc, J.-P, J. M. Roucny, M. FerRANDINI, AND J. FERRANDINI. 2007. Editorial: The Messinian Salinity Crisis revisited. [bis]. Geobios, 40: 231-232. 58 FIELDIANA: ZOOLOGY _— 4 a ona . ie - Row Pa i cP ; aa. “sie © ee ——— a ; iS trade we etal ian £2 ree ee Amaae a v, 4 ones ae =: 3 ~ , ( eee | hd moh, ‘ a VAS ae ova até a F — ° : a. i ¥ g «¢ % aa = " ~ 7 ron > eee * al a ~ =A ~~ ¢ f 7 ; 7% ~ 5 7 . * 7 a ad ' : a . 77> : if J v4 e. = “Ts, : 7 9 a : e c a : 7 ’ { ay, ’ ‘ 7 < . P . > F Mf | / ; ‘/. = .. : i, 7 » ee -—= _ a ' aE ¢ rs eel ye * ie ” lo i Se eer at a ‘ee ct. va, UR gui t - . =. © wt i Tonk! Ae at Bur. Micitcen: . Pash fs tad eee HIN Nee Aa s eI uh We ve . *, tee , A sie eet ap « . ( $ DA Ia aade, ACO aC aes PON Hg : a > f hs