sign in sign up
<<
Home , Assam macaque, Barbary macaque, Blyttia (plant), Cercopithecinae, Ennedi Plateau, Grivet

JASs Invited Reviews Journal of Anthropological Sciences Vol. 87 (2009), pp. 33-91

The of the western Palaearctic: a biogeographical, historical, and archaeozoological review

Marco Masseti1 & Emiliano Bruner2

1) Dipartimento di Biologia Evoluzionistica “Leo Pardi”, Università di Firenze (Italia) e-mail: marco.masseti@unifi .it 2) Centro Nacional de Investigación sobre la Evolución Humana (CENIEH), Burgos (España) e-mail: [email protected]

Summary – e Western Palaearctic is traditionally regarded as a zoogeographical unit which is lacking in primatological fauna. e representatives of this taxonomic group which has been documented within its boundary can be referred to the genera Macaca, Papio, and , and possibly also to and . e data for the present research were collected through a review of all previous knowledge of the primates of this biogeographical , including their history, and through original sightings and direct observation of fi eld signs. Surveys were carried out directly in North , the peninsula of , and in the . Additional data on distribution were obtained through the examination and evaluation of the materials conserved in several museums. A historical and archaeological investigation was also carried out, appraising both archaeozoological fi ndings and prehistoric and ancient artistic production, in order to evaluate the importance of the monkeys of the Western Palaearctic in relation to local activities and needs.

Keywords – Ethnozoology, Macaca, Papio, Chlorocebus.

“Hamadryads. A type of -nymph. of north of the Himalaya along northern All dryads are concerned with and live in Africa, including the northernmost part of the them when they are roaming the Sahara (cf. Corbet, 1978). Like all faunal , - but each hamadryad is the spirit of a however, there can be no precise defi nition of particular . the Palaearctic (Vaurie, 1965; Cramp, 1977). In She lives always in this tree and speaks out to fact, zoologists have frequently found diffi culty in warn woodman away delimiting this zoogeographical area. Ellermann when they try to set an axe to her tree. & Morrison-Scott (1951), for example, argued When the tree dies, she dies too” that certain arbitrary limits must be set in its defi - (Evslin, 1988) nition. ey suggest drawing the African bound- ary along the 20°N parallel which, considering the barrier of the Sahara: “… does correspond reason- The Western Palaearctic ably well with the facts”. Nevertheless, this means that several Saharan mountainous complexes and e Palaearctic region has been recognised and their peculiar biocenoses, such as the Nigerian Aïr acknowledged as a natural zoogeographic region massif, and archipelagos, such as Cape Verde and/ since Sclater fi rst proposed it as far back as 1858. or the Farasan Islands (), are not com- It can be defi ned approximately as the prised within the borders of this zoogeographical

the JASs is published by the Istituto Italiano di Antropologia www.isita-org.com 34 The primates of the western Palaearctic

range, which does however include areas such & Chiu, 1983). Its diff usion to the north may as the Tassili n’Ajjer (), and the Tibesti not extend beyond 30°26’N, 102°52’E (Groves, (). Corbet (1978) instead suggests that the 2005). However, broadly speaking, the northern- African boundary of the Palaearctic Region starts most limit of the dispersion of primates in the in the west at 21°30’N, between Rio de Oro and Palaearctic is represented by the natural diff usion , continues across at the same lati- of the Japanese (apart from the “every- tude and thereafter follows the political boundaries where” called sapiens). is comprises so including Algeria, and entirely and most of the Japanese archipelago, including even excluding the whole of , Chad and . the southern areas of the island of Hokkaido, e entire is included, as are located above the 42°N parallel, at the north end the Ahaggar mountains (Algeria), while most of of opposite the Russian island of Sakhalin. the Tibesti range is excluded. Furthermore, in e northern pig-tailed macaque, Macaca leonina detailing the current western boundary of the (Blyth, 1863), is distributed as far as Palaearctic Region, Corbet (1978) includes the (), north of 25°N, whereas the distribu- archipelagos of Spitzbergen, Iceland, the Azores, tion of the macaque, Macaca assamensis Madeira, and the Canaries, but excludes the (McClelland, 1839), ranges from the Himalaya islands of Cape Verde. On the other hand, follow- foothills up to 2,750 m. in central east ing Vaurie (1959-1965), Cramp (1977) suggests into and southeast China (Groves, 2001). comprising within the Western Palaearctic all the e Formosan rock macaque, Macaca cyclotis eastern Atlantic islands south of the Cape Verde (Swinhoe, 1862), may also fall within the bound- archipelago, adding the Banc d’Arguin group ary of the Palaearctic, being naturally limited to the (Mauritania) - but not the adjoining Mauritanian island of Taiwan, comprised between 22°N and mainland - where the extensive research of de 26°N (Groves, 2001). Corbet (1978), however, Naurois (1969 and 1994) have clearly demon- includes only a handful of primates among the strated the Palaearctic character of the avifauna. faunal elements of the Palaearctic Zoogeographic Although the easternmost territories of the Indian Region: 4 and 1 . In fact, for the subcontinent, such as Gujarat and Rajasthan, Macaca, he considers the two M. are not biogeographically comprised within the sylvanus (L., 1758), and M. fuscata, respectively boundary of the Palaearctic, they nevertheless isolated in and in the Japanese archi- display several zoogeographical elements char- pelago, together with M. mulatta (Zimmermann, acteristic of the latter biogeographical region (cf. 1780), native of central , from to Masseti, 2002b); south of the Sahara, the moun- southern China, and probably introduced in an tainous slopes of are characterised by area in the surroundings of Beijing (Ellermann & the relic distribution ranges of the ibex, Capra ibex Morrison-Scott, 1951) but apparently extirpated L., 1758, and other Palaearctic taxa. in 1987 (Zhang et al., 1989; Fooden, 2000); Although not as numerous as those distrib- and M. cyclopis, native of Taiwan, as mentioned uted in the Afrotropical, Oriental and Neotropical above, and imported onto the Japanese islands biogeographical regions, the Palaearctic monkeys of Oshima and Izu. e only representative of comprise various species, including inter alia the the genus Papio, considered by Corbet (1978), , Macaca sylvanus (L., 1758), is the hamadryas, Papio hamadryas (l., 1758), a the , Macaca fuscata (Blyth, species still dispersed in eastern Africa and in the 1875), and possibly the snub-nosed , southern Arabian peninsula, which he reputed to Rhinopithecus roxellana (Milne-Edwards, 1870). have inhabited northern Africa and the Egyptian is monkey is the leaf-eating primate with the territories in ancient times. Some years earlier, most northerly distribution, occurring in the Ellermann & Morrison-Scott (1951) also consid- mountainous areas of Sichuan and the south- ered the sacred baboon as the only representative ern parts of Gansu, Hubei and Shaanxi (cf. Ho of the genus occurring in the Palaearctic region M. Masseti & E. Bruner 35

due to the fact that they took the African south- the boundary of the Western Palaearctic up to ern boundary of this biogeographical region to be the border with the eastern Indian sub-continent. the 20th parallel. Papionins are well documented Furthermore, Martin & Hirschfeld (1998) pro- in the Western ethnological environments, hav- pose the inclusion of the entire Arabian peninsula, ing represented an attractive topic for painters excluding two small areas where Afrotropical infl u- and other artists from the earliest to more recent ence is dominant - a land strip along the south- historical times (Plate 1). ern coast of the , and the Afrotropical e Western Palaearctic is the western por- enclave between Dhofar () and the Mahral tion of the Palaearctic zoogeographic region Province of – together with part of (cf. Cramp, 1977). e suggested boundaries in the geographical area formally treated as the exclude Greenland (Cramp, 1977), but include western subdivision of the Palaearctic. Here, the all the islands of the eastern Atlantic Ocean up primate belt extends from approximately 36°N in to Cape Verde, i.e. the whole of Macaronesia, the northern to approximately 20°S in including the Azores, Madeira and the Canary the southern Sahara which, as mentioned above, islands (Vaurie, 1959; Coutinho Saraviva, 1961; is roughly considered the southern boundary of Naurois, 1969; Sunding, 1970, 1979; Kunkel, this biogeographical unit. To the north, monkeys 1980; Gonzales Henriquez et al., 1986; Beyhl et are not found outside this geographic limit, with al., 1995). us the northernmost boundary of the exception of small communities introduced by the Western Palaearctic would be located in the , such the M. sylvanus colony of Gibraltar north-eastern Atlantic Ocean and in Sea. (cf. Napier & Napier, 1985) (Fig. 1). Vaurie (1959-1965) and Cramp (1977) include is current research is the result of a series the Sahara south of the northern borders of the of studies carried out in several of the territories region within the perimeter of this zoogeo- of diff usion of the Western Palaearctic mon- graphical area. However, they include the moun- keys. Data were collected through a review of tain massif of Tibesti, which is located above the all previous knowledge of the primates of this 20th parallel and exclude those of Aïr (Niger) and zoogeographical region, including their his- , where the Afrotropical element predomi- tory, through original sightings and direct nates. To the east, the boundary could be limited observation. Surveys were carried out directly to the northern Arabian peninsula (cf. Harrison, in Northern Africa, the peninsula of Gibraltar 1964), or alternatively could run northwards (UK), and in the Sahara. Additional data on the along the eastern border of (cf. primate distribution were also obtained through Tutin et al., 1964), and of the , includ- the examination and evaluation of the materi- ing almost all of Iran (Guest & Al-Rawi, 1966; als conserved in the following museums: the Masseti, 2002). In any case, as Cramp (1977) Forschungsinstitut and Natural History Museum argues, if the boundary of the Palaearctic region Senckenberg, Frankfurt am Main (SMF); the eludes any logically unequivocal demarcation, the Natural History Museum in London (BMNH); determination of the eastern limits of its western the Odontological Museum of the Royal College part is clearly largely arbitrary. us, the same of Surgeons of London (RCS OM); the Institut author includes all European Russia, using the Français d’Afrique Noire, Dakar (CG. IFAN); limits which are internationally accepted for the the Libyan Museum of Natural History of the Flora Europea (Tutin et al., 1964), and, further Assaray Al-hamra Museums, Tripoli; the Natural south, he includes the remaining regions of the History Museum of the University of Florence, former USSR between the Caspian Sea and the Zoological Section “La Specola” (MZUF); and Black Sea, , Iraq and Kuwait, but excludes the “Giuseppe Sergi” Museum of Anthropology the whole of Iran. However, according to other of the “La Sapienza” University of Rome. A his- authors, such as Guest & Al-Rawi (1966) and torical and archaeological investigation was also Masseti (2002), Iran is wholly comprised within carried out, appraising both archaeozoological

www.isita-org.com 36 The primates of the western Palaearctic

fi ndings and prehistoric and ancient artistic and the , Chlorocebus sabeus (L., productions, to evaluate the importance of the 1766), to which we may possibly also add the monkeys of the Western Palaearctic in relation , Chlorocebus aethiops (L., 1758). Another to local human activities and needs. Original representative of the Order of Primates (Family archaeozoological data from the Tadrart Acacus Galagidae), the bushbaby, Galago senega- (Libyan Sahara), and one modern osteological lensis É. Geoff roy, 1796, has also been recorded fi nd from the archipelago of Farasan (southern from one of the areas adjacent to the southern Arabia), are also discussed. An additional aim of boundary of the Western Palaearctic, the Saharan this work is to attempt to concentrate the entire mountainous massif of Ennedi, in Chad. existing bibliography regarding the primates of Individuals of several of these species have often the Western Palaearctic. been transferred beyond their natural distribu- tion for various human needs. It is not immedi- List of the species ately apparent why humans should have wanted Within the vast geographic area represented to export these , and the phenomenon by the Western Palaearctic, there are very few can only be explained by considering each case taxa of primates which existed – and still exist– individually. However, evidence suggests that in as wild or feral free-ranging populations. ere many cases they were transferred voluntarily by are essentially six or seven species: the Barbary humans. Furthermore, historical, ethnozoologi- macaque, Macaca sylvanus (L., 1758), the olive cal and archaeozoological studies document the baboon, Papio anubis (Lesson, 1827), the sacred fact that monkeys were utilised as food, for medi- baboon, Papio hamadryas (l., 1758), the patas cine, as pets and for other purposes from prehis- monkey, Erythrocebus patas (Schreber, 1775), toric times onwards.

Fig. 1 - Boundaries of the Western Palaearctic within the palaeontological (grey) and extant (dark grey) range of non-human primates. The suggested boundaries of the Western Palaearctic com- prise within their perimeter the Sahara south of the northern borders of the Sahel region, including the mountain massif of Tibesti, which is located above the 20th parallel, and the archipelago of Cape Verde. M. Masseti & E. Bruner 37

Barbary macaque or Barbary ape, According to Szalay & Delson (1979), this Macaca sylanus (L., 1758) genus appears to be of North African provenance, where it seems to have developed in the Miocene French: magot and is still present. e mtDNA evidence con- Spanish: mona de Gibraltar, mona de Berberia fi rms the deduction from morphological studies, Italian: bertuccia, bertuccia berbera corroborating hypotheses of the Africa origin for German: magot the genus Macaca (Morales & Maelnick, 1998; Fooden, 2005). Two or three hundreds years ago A unique characteristic of Macaca sylvanus the monkeys were already living in the southern (L., 1758) is the absence of a tail, as in true (Richard et al., 1989), where . is is the only macaque found outside they are present today on the Asia (Richard et al., 1989). Linnaeus described (UK) (Fig. 2). e present distribution of the its in his (10th ed. species is the remnant of a much more wide- 1: 25), on the basis of the examination of speci- spread population which, up to the last glacial mens from the “”. A portrait of episode, inhabited vast areas in and Barbary macaque already fi gured among the Africa (Camperio Ciani, 1986). In fact, dur- various representations of animals in Gesner’s ing the M. sylvanus was widespread Historia Animalium, the fi rst edition of which not only in northern Africa but also in Europe was printed in Zurich in 1555 (Plate 2a). (Kurtén, 1968; Delson, 1980; Camperio Ciani, Nevertheless, it is not possible to determine the 1986; Kowalski & Rzebik-Kowalska, 1991). terra typica of this species which Linnaeus had Older African fi nds date back to the Pliocene established on the basis of the very approxi- (Ain Brimba, Ichkeul) onwards (Kowalski & mate representation of a captive specimen, pub- Rzebik-Kowalska, 1991). Neolithic remains of lished by Prosper Alpin in his Historia Naturalis the species are known from the littoral Algerian Aegypti (1581-1584), vol. II, pl. XVI (cf. site of Boulevard Bru (Flamand, 1902), and Cabrera, 1914; Fenoyl, 1980). (Plate 2b). e the upper part of Djebel aya, in the Saharan Barbary macaque is the only primate indigenous Atlas (Bourguignat, 1870). Also Gautier (1993) to the Western Palaearctic, where it occurs - comprises this monkey in the archaeozoological ther north than any other non-human species, inventory of the Maghreb. Barbary macaques also being the only macaque found in Africa are extinct in and Libya (Haltenorth & (Waters et al., 2007). Together with the wild Diller, 1977). , Oryctolagus cuniculus (L., 1758), several In their extant Maghrebi distribution, the representatives of the genus Lepus, the Iberian macaques are most abundant in high, mixed lynx, Lynx pardinus (Temminck, 1827), and a cedar and oak forest (Deag, 1977; few other species, M. sylvanus fi gures among the Fa, 1984; Ménard et al., 1985; Macharias et al., medium sized endemic to the west- 1999). ey can be encountered in the upper ern Mediterranean zoogeographical unit (cf. reaches of the mountains in winter, despite low Masseti, 2002). Barbary macaques are confi ned temperatures and snow (Kowalski & Rzebik- to fragmented populations throughout their Kowalska, 1991). At lower elevations, decidu- current natural range, western North Africa, ous and evergreen oaks are dominant, where they are discontinuously distributed in pressure is intense, cultivation widespread, and the subtropical mountainous areas of monkeys occur only in certain , always at and Algeria (north-western Maghreb) (Cabrera, low density (Deag, 1977; Fa, 1984; Macharias 1932; Richard et al., 1989; Fa, 1999). ey are et al., 1999). ey are also found at very low separated from the nearest Asiatic population of density in scrub forest and on treeless mountain the genus Macaca by a distance of more than ridges. Evergreen forests of cedar and oak appear 5000 km (Camperio Ciani, 1986). to be the optimal biotope, where the population

www.isita-org.com 38 The primates of the western Palaearctic

density is highest and there is also the highest the Moroccan populations of these macaques survival rate of youngs (Ménard et al., 1985). In are very fragmented (Cuzin, 2003). In northern the forest these parameters are lower. Morocco, a few scattered groups can be found Persecuted in the plains because of damage to in disturbed in the and the coastal crops, and gardens, they are partial to Mediterranean region (Lakhdar et al., 1975; Fa rocky areas, featuring gorges and rock shelters. et al., 1984; Mehlman, 1989). Macaques are cur- Rocky slopes without arboreal , how- rently reported from four areas of the Djebela ever, are unfavourable for these primates, because region: Djebel Moussa, west of the Spanish pos- they always need to be near water. Over its entire session of Ceuta; El Haouz, south of Tetouan; range in Morocco and Algeria, the species was Djebel Bou Hassim, south-west of Tetouan; and previously estimated to number 9,000-23,000 Djebel Talassamtane, south of Chaouen (Waters individuals (Fa et al., 1984; Oates, 1996), but et al., 2007). Recent surveys indicate that the recent population estimates in the species’ strong- total population of the Djebela has dropped to hold, the mountains in Morocco, no more than 200-300 individuals. Although record a dramatic decrease in number over the some areas where the species is found are now last decade (Camperio Ciani et al., 2005). is protected, further commitment to safeguarding population decline is attributed to the loss the species will be crucial in northern Morocco. of prime , mainly cedar forest, which In the southern part of the same country, a small has signifi cantly decreased due to the growing relic population also dwells in the western High impact of overgrazing by mixed fl ocks of goats (Fa et al., 1984; Mehlman, 1989; and sheep, and the consequent degradation of Cuzin, 2003), being present in the valley of the the forest. Human-caused habitat Oued Ourika, south-east of Marrakesh. In this in the Middle Atlas further risks compromising area, monkeys have been recorded from the low the future of the world’s only remaining large M. gorges of Assif Tinzer up to the western slopes sylvanus population. Outside the Middle Atlas, of Jbel Yagour, and also in the high valley of

Fig. 2 - The dispersion of Barbary macaques is today confi ned to western North Africa (Morocco and Algeria), and the Rock of Gibraltar, in south- (photo Emiliano Bruner). M. Masseti & E. Bruner 39

Oued Zat (Deag & Crook, 1971; Cuzin, 1996). in the Roman world as novelties and curiosities Around the mid 1980s, the population of the (King, 2002). Pliny the Elder made various com- Ourika valley was estimated at about one hun- ments on monkeys in general (Naturalis Historia, dred individuals (Aulagnier & evenot, 1986). 8.215-216), particularly on their intelligence and According to Cuzin (2003), the continuous cunning, but he only distinguished tailed from decrease in number of these monkeys indicates tailless primates in a generic manner. Apes and the vulnerability of the population. monkeys were often dressed up as performers, In Algeria the species is dispersed throughout with various articles as props (McDermott, 1938; the regions of Djurdjura and Kabilia (Kowalski Toynbee, 1973), and this practice appears to have & Rzebik-Kowalska, 1991; Fa, 1999), where continued up to Medieval, Renaissance, and even it occurs from sea level up to the mountain modern times. An illustration of this can be found tops (the highest peak of the Djurdjura Mts. is in a detail of the mosaics that decorate the north- 2,308 m a.s.l.). Specimens from the “Gorge de eastern hall of the imperial palace of Byzantium la Chiff a” (Gargide Chiffa), 36°30’N 2°45’E, are (register I, sector C, panel Co), Istanbul (Turkey), present in the collection of the Natural History where a clothed monkey is attempting to catch Museum of London (BMNH 1939.3471), and a bird from the top of a tall tree, using a lime- the University Museum of Zoology of Cambridge twig (Jobst et al., 1997) (Plate 3). e production (CMZ E7495A, E7495B). e former institu- of these mosaics dates to the fi rst half of the 6th tion also holds another Algerian specimen from century AD. Eff ectively, the Roman portrayal of Stora (BMNH 49b). According to Kowalski & monkeys is largely derived from Eastern, Greek, Rzebik-Kowalska (1991), some of the Algerian and Hellenistic art, as demonstrated by the pre- territories still inhabited by Barbary macaques ponderance of catalogue entries from those sources were probably more extensive in the past, and the in McDermott (1938). Barbary monkeys, in par- animals also occurred in other areas. Nevertheless, ticular, were probably more accessible to Roman the same authors are of the opinion that the range hunters and trappers than other species, but this of these primates has not been greatly reduced. does not rule out the importation of specimens Genetic research indicates that the world popu- of other primates from more far-fl ung areas, such lation of the Barbary macaque is divided into at as sub-Saharan Africa and/or the Indian subconti- least 5 diff erent subpopulations (von Segesser nent. Among the ruins of Pompeii (Naples, south- et al., 1995; Martin & von Segesser, 1996; von ern Italy), osteological and artistic evidence of the Segesser et al., 1999). artifi cial presence of the monkey is not rare (King, 2002). For example, the incomplete postcranial Barbary macaques in antiquity and the Middle Ages remains of a unique fi nd of a juvenile Barbary Together with the baboon and the , macaque skeleton was identifi ed using osteology the Barbary macaque was perhaps the best-known and ancient DNA tecniques, although its prov- species of primate in ancient times. Despite it enance is unspecifi ed (inv. no. Lab. 16) (Ciarallo seems that there are no depiction of these mon- & De Carolis, 1999; Bailey et al., 1999). It is key in Pharaonic art (cf. Groves, 2006), at least 20 most likely to have been an exotic pet, probably a skulls of M. sylvanus were instead provided by the macaque, and therefore, according to King (2002), archaeological investigation of the sacred probably, a Barbary ape. In England, osteological necropolis at Saqqara, dating from after 300 B.C. remains (maxilla and palate, calvarium) of a sub- (Goudsmith & Brandon-Jones, 1999). It was the adult M. sylvanus were provided by the archaeo- pithekos of Aristotle, being more widely distrib- logical exploration of Catterick Fort (Yorkshire, uted during classical times, thus generating a gen- UK) chronologically dating to the Romano- eral portrait of the tailless “apes” of Africa and Asia British period (cf. Napier, 1981; Lynn, 1997). (Spencer, 1995; Rolfe & Grigson, 2006). It seems ey are now preserved in the collection of the that monkeys of various sorts were not uncommon British Museum, London (BMNH 1977.3120).

www.isita-org.com 40 The primates of the western Palaearctic

Lynn (1997) recorded other “British” remains of position have made it possible to date the inhu- the species from Dunstable, Bedforshire, dated to mation of the monkey to between the 5th and the late 2nd century A.D. 6th centuries AD. e military characteristics of However perhaps one of the most intriguing its apparel suggest that the Barbary macaque may fi nds is that of an exemplar which came to light have been either a pet or a military mascot. in , in the old Roman city of Iulia Livica M. sylvanus was perhaps the most famous (Llívia, Cerdanya), in the vicinity of Empúries primate in the Western World even in medieval (Catalonia), where the tomb of a Barbary macaque times (Wendt, 1959; Masseti, 1991). In 1558, was found (Guardia & Maragall, 2004; Aliqué, an engraving representing a monkey hunt was 2007) (Fig. 3). Discovered along with the buried comprised in an extensive series of plates depict- animal was a series of military decorations that ing various aspects of the hunt in sixteenth cen- were part of its personal accoutrements, including tury Europe (Plate 4). is was published in belt buckles and a number of bronze plaquettes Antwerp, with the title of Venationes Ferarum, (Guardia et al., 2005). Both the chronology Avium, Piscium, by the Flemish artist Jan van der attributed to these fi nds and their stratigraphic Straedt, also called Antonio Stradano (cf. Baroni Vannucci, 1997). Several peculiar characteristics of the morphological rendering of the monkeys, and in particular the complete absence of any tail suggest that the animals portrayed are undoubt- ably M. sylvanus. Via the Mediterranean, the Barbary macaque spread extensively to Italy (Hill, 1966), where it has frequently been portrayed in artistic artefacts from at least the end of the 14th century. It is in fact included, along with reproductions of other animals, in the famous Giovannino de’ Grassi sketchbook preserved in the Biblioteca Civica Angelo May of Bergamo (Codice Cassaf. 1.21, f. 5r.) (cf. Recanati, 2005). Barbary macaques are presented to the new- born Christ in the panel of the Adoration of the Magi, painted by Gentile da Fabriano between 1420 and 1423 (Florence, Uffi zi), and in the fresco by Benozzo Gozzoli on the same subject in the Florentine Palazzo Medici Riccardi. In Renaissance Florence, the image of the species is evoked in various art works in the gardens of Boboli. Several bronze Barbary macaques decorate the Fontana delle Scimmie (“fountain of the mon- keys”), formerly erroneously ascribed to Pietro Tacca, and possibly inspired by the bronze sculp- tures of primates created by Giambologna for the Florentine fountain of “Samson and the Philistine” Fig. 3 - Drawing of the tomb of a Barbary macaque from the ancient Roman necropolis in the Casino of San Marco (Masseti 1991). e of Iulia Livica (5th-6th centuries AD, Catalonia). images of two almost life-size Barbary macaques, The animal was buried with military decorations in polychrome marble (marmo mistio), were also (belt buckles and bronze plaquettes), suggest- realized by the sculptor Cosimo Fancelli - pos- ing it may have been a military mascot (after Guardia et al., 2005). sibly to a model by Baccio Bandinelli - around M. Masseti & E. Bruner 41

1555, in the Grotta degli Animali or Grotta del are known to have resided there for at least the diluvio of the Medici Villa of Castello, near past 265 years. Groves (2001) refers to a pos- Florence (cf. Masseti, 1991; Acidini Luchinat, sible introduction of the species in Roman times. 1992; Paolucci, 2000; Masseti, 2008) (Plate 5). Others sustain that the animals may have been However, one of the most amazing representa- introduced onto the Rock from North Africa by tions of this primate of all Italian art is again the Moors who, under the Saracen Tariq, cap- to be found within the perimeter of the Boboli tured and fortifi ed Gibraltar in 710-711 AD gardens: in the frescoes painted in the fi rst hall (Hill. 1966; Lever, 1985). Mediaeval chronolo- of Buontalenti’s Grotta Grande by Bernardino gies also coincide with the earliest appearance Poccetti between 1586 and 1587, during the last in the Iberian peninsula - and in Europe - of years of the Grand Duke Francesco I de’Medici, other African mammals of medium size, such (cf. Chiarini, 1977). One individual of Barbary as the common genet, Genetta genetta (1758) macaque is portrayed sniffi ng a Damascene rose (cf. Morales, 1994), and the Algerian hedgehog (Plate 6). is was the same attitude in which (Morales & Rofes, 2008). However, since no Suleyman the Magnifi cent, extraordinary sultan mention of monkeys is made during the period of the Ottoman empire from 1520 and 1566, of Spanish occupancy between 1492 and 1704, generally chose to be portrayed. Suleyman, who it is possible that the Moorish importation died doubled the territory of the empire during the out, subsequently being replaced by new stock 46 years of his rule, wished to be painted in this (cf. Lever, 1985). On the other hand, Zeuner pose which emphasised his sophistication and (1952) considers that “… it is certain that apes sensitivity rather than underscoring his fame as were present on the Rock when Gibraltar was cap- a ruthless conqueror. e domain of Suleyman tured by the British in 1704”. Following Cabrera the Magnifi cent extended from the surround- (1914), Garcia (1979) and Castells & Mayo ings of Vienna far to the east and into Egypt and (1993) are also of the opinion that the macaques Persia, while his fl eet dominated the Red Sea and were already present on the Rock before the virtually the whole of the . British occupation, whereas Fa (1981 and 1999) Consequently, it’s not diffi cult to imagine why declares that they were released on the prom- the artist who evoked the scene of the “sniff - ontory in the early 1740s. In eff ect, the earliest ing monkey” in the Boboli gardens was asked certain account of monkeys on Gibraltar dates by Francesco I, or someone else at the Medici back only to this year, when mention is made court, to paint a caricature of the Muslim enemy. of a large importation of apes “and other game e image of a Barbary macaque was thus trans- from Barbary”, and when a poll tax was levied formed into an amazing personifi cation of the on “apes, Moors, Jews and other aliens”( cf. Lever, detested Islamic world and its ruler. 1985). At this time, on Gibraltar, monkeys were regarded as game for hunting, for the enhance- Macaques on the Rock: the colony of Gibraltar ment of which other animals including Barbary As mentioned above, the present distribution partridges, Alectoris barbara Bonaterre, 1790, of M. sylvanus comprises a European enclave coin- were also imported onto the Rock from the ciding with the boundary of the promontory of Maghreb (cf. Lord Lilford, 1866; Cortés et al., Gibraltar, a British possession in the Iberian pen- 1980; Johnsgard, 1988). It was in fact not until a insula since 1704 (Zeuner, 1952). e Barbary century later, in 1856, that a Garrison Order was apes of Gibraltar constitute the oldest established fi nally issued which prohibited the killing of the colony of free-ranging monkeys in Europe (Lever, monkeys for any reason. Despite this, seven years 1985). In fact, the anthropochorous origin of later only three macaques remained on the Rock this population is beyond all doubt. Although (Lever, 1985). From this time on, Barbary apes it is uncertain exactly when the macaques were have always been protected within the perim- fi rst introduced to the Rock (Hill. 1966), they eter of Gibraltar and their contingents restocked

www.isita-org.com 42 The primates of the western Palaearctic

from North Africa, as need arose. Files kept at the or Anubis baboon, Gibraltar Regiment tell the story of the “Rock Papio anubis (Lesson, 1827) apes” from 1913, the year when a Master Gunner was fi rst given the task of feeding the macaques French: babouin doguera, papion anubis (Garcia, 1979). ere are reports of at least three Spanish: papión perruno other introductions of monkeys from Morocco to Italian: babbuino verde Gibraltar, the last of which (1939-1943) gave rise German: Anubispavian or Steppenpavian to the current population of Gibraltar (Anon., 1880; Garcia, 1979; Fa, 1999). It was tradition- Today, are unknown in North ally believed that the Gibraltar stock essentially Africa, occurring only from the central Sahara originated from those monkeys found in Anjera southwards, their northernmost limit of dis- on the Gibraltar Straits, in the vicinity of the persion being around 15°N (cf. Haltenorth & aforementioned Jebel Mûsa (Djebel Moussa), Diller, 1977). From a taxonomic point of view, “… known to Europeans as Ape’s Hill” (Meankin, fi ve species are currently recognised within the 1901). Recent genetic analyses eff ectively con- genus Papio, (Muller, 1773). Among them, the fi rm that the extant population of macaque in olive baboon, Papio anubis (Lesson, 1827), is a Gibraltar descended from a handful of individu- large sized baboon which is still dispersed from als imported during World War II from north- Mali to (Zinner et al., 2001), western Africa. ey were, however, found to and north-western (Napier & Napier, include Algerian and Moroccan haplotypes, 1967; Groves, 2005). is is an Afrotropical spe- revealing a dual origin of the founding females cies which could have been represented by some (Modolo et al., 2005). Other genetic studies populations in ancient north-eastern Africa, and show that human intervention has had marked possibly in Egypt. eff ects on genetic equilibrium and heterozygosity (Segesser et al., 1995; Martin & Segesser, 1996; The olive baboon in ancient Egypt Segesser et al., 1999). It is in fact possible to trace As with other primates, in Egypt there are the origin of the founders of the Gibralter colony, no early osteological records of distribution, back to Morocco, probably to the Middle Atlas, although baboons have frequently been found as perhaps with some residual infl uence of animals mummies, even in great numbers (Kessler, 1989; imported earlier from Algeria. Osborn & Osbornová, 1999). Boessneck (1988), Macaques are traditionally confi ned to the for example, recorded that mummies of this spe- higher and less frequented parts of the Rock of cies had been found in the Tuna el Gebel necrop- Gibraltar (cf. Sclater, 1900). At present, there olis, west of Hermopolis. Olive baboons were are some 300 animals in fi ve troops occupying also found at ebes (Osborn & Osbornová, the area of the Upper Rock, with occasional for- 1999). Various other sites that yielded Late ays into the town in monkey mayhem. Despite Period (1085/1070-332 BC) mummies of the this, the macaques are a tourist attraction. species include Hermopolis Parva (Damanhur, Notwithstanding extensive evidence of viral in the western Delta), Tanis (Zoan, in the east- transmission from macaques to humans associ- ern Delta), Kom Madinet Gurab near El Lahun ated with biological wet vectors (urine, blood, southeast of El Fayum, Mustai in the south cen- saliva – see Brack, 1987; Cavicchio & Friedrich, tral Delta, and some questionable remains at 2006), the monkeys are allowed free contact with Saqqara (Osborn & Osbornová, 1999). In the visitors. Viral infections are highly dangerous and art of the Dynastic period, illustrations of Anubis can even lead to death (e.g. through meningitis). baboons are rare. e phenotypic characteristics In view of the lengthy course of such patholo- of the species, such as the protruding nostrils and gies, the correlation with the original source of angled tail, are not clearly represented, nor can contamination is not easily recognisable. facial and buttock coloration always be verifi ed. M. Masseti & E. Bruner 43

e artistic evocation of olive baboons appears which this extreme northern population has – on one of the ships being loaded in the Punt undergone gives it great biological signifi cance. expedition launched by the Queen Hatshepsut, e feeding habits of the Aïr olive baboons are in together with the homecoming ships laden with fact mainly based on the consumption of dates bounty from Punt - in the mural decoration of from the dom palm, Hyphaene thebaica (L.), and, her temple at Deir el Bahari, in ebes (18th to a lesser extent, of sp. and other Dynasty, around the middle of the 2nd millen- (Bousquet, 1992). In this regard, an inter- nium BC). One of the best representations of esting fact emerging from Keimer’s (1939) dis- these baboons is, however, reputed to come from cussion of the baboon and the dom palm is that the tomb of Khnumhotep, of the 12th Dynasty the of the tree were called “nuts” in ancient (1991-1785 BC), at Beni Hasan (Osborn & Egypt and was considered as a nourishing food. Osbornová, 1999). e animals are depicted Its importance is emphasised by numerous artis- in a fi g tree, and are a dark green colour with tic representations showing baboons with sacks reddish-brown faces and callosities and without of dom fruit and climbing trees (cf. Osborn & mantles (Plate 7). Ermann (1894) regarded them Osbornová, 1999). as monkeys helping with the harvest. Other In their northernmost range of distribution, authors, such as Houlihan (1997), consider that olive baboons extended beyond the southern this is unlikely, since the baboons appear to be borders of the Western Palaearctic, having been about to eat the fi gs in their hands. Goudsmith formerly reported from Tibesti. is is an iso- & Brandon-Jones (1999) identifi ed 149 skulls of lated mountainous massif, about 3,500 metres in P. anubis in the baboon catacomb of the sacred height (Petragnani, 1928; Brown, 1965), which animal necropolis at Saqqara (after 300 B.C.). is located in the middle of the Sahara desert In ancient Egyptian art, the distinction between between about 19°N and 24°N. Due to its geo- olive baboons and other representatives of the graphical position, the entire region falls within same genus, i.e. Papio hamadryas (L., 1758), is the confi nes of the western Palaearctic, together made possible mainly through an evaluation of with the rest of the Sahara south of the northern the typical rendering of the angled tail, as well as borders of the Sahel area (cf. Vaurie, 1959-1965; the coat colour and the lack of the mantle. Cramp, 1977). In the early 1950s, a subspecies of olive baboon was described from Tibesti as Saharan baboons P. anubis tibestanus Dekeyser & Derivot, 1960 e olive baboon also includes some popu- (Dekeyser, 1952). lations in the southern mountainous massifs of the Sahara (Le Berre, 1990), from where it has Tibesti baboon, Papio anubis tibestianus Dekeyser been reported in the remote mountains of Aïr & Derivot, 1960 (Niger), Tibesti (Chad), Manakaoki ( omas, Western scientists were not informed of the 1925), or Wadaï and Ennedi (Chad) (Bourbon, existence of a population of these monkeys in the 1932; Dalloni, 1935; Dekeyser, 1950 and 1952; mountainous area of Tibesti until around the end Dekeyser & Derivot, 1959; Hill, 1966; Gillet, of the 19th century (Lavauden, 1926; Petragnani, 1968; Haltenorth & Diller, 1977; Le Berre, 1990; 1928; Dalloni, 1935; Dekeyser, 1952 and 1955; Bousquet, 1992). More specifi cally, according to Dekeyser & Derivot, 1959; Beck & Huard, Malbrant (in Dekeyser, 1955), in Chad baboons 1969). e history of baboons in this region is still are not found beyond the northern limit of dif- far from being well known. According to Hufnagl fusion represented by the 13th parallel, while (1972), the German explorer Gustav Nachtigal they extend eastwards as far as Tibesti through was the fi rst scientist to reach the mountains of the Ouadaï and the Ennedi. In Aïr, the species Tibesti around 1870, and he nearly died in the only lives in the massif of Tamgak (wadies and attempt. He reported the existence of monkeys mountains) (cf. Hall et al., 1971). e isolation there in the extreme south of Libya (Hufnagl,

www.isita-org.com 44 The primates of the western Palaearctic

1972). Later on, little by little, more information located between Zouarkè and Kachem. is spec- was gathered, and in 1926 Lavauden observed imen had already been presented to the Première that in the region of Tibesti: “… Colonel Tilho Conférence Internationale des Africanistes de assured us that there were cynocephali there”, pos- l’Ouest in Dakar in 1945 by J. Bigourdan (1950), sibly attesting the occurrence of baboons in this but was only later described by Dekeyser (1952). In Saharan region for the fi rst time. A few years later, 1957 two young French explorers, Carl and Petit Petragnani (1928) again referred to the occur- (1954) collected specimens of the Tibesti baboon rence of these primates in Tibesti, and in particu- in the western range of the Saharan mountainous lar in the vicinity of the town of Zouar, located massif, again near Zouar, and in 1960 Dekeyser in the foothills of the western slopes, along the and Derivot described the new material as a short- valley of “Enneri Durso”. In 1952 Dekeyser iden- tailed subspecies of the olive baboon, i.e. P. anubis tifi ed the only skull of baboon then known from tibestianus. A skull of this rare taxon was given by Tibesti (cf. Hufnagl, 1972). Its type specimen was Mr. Kenneth Guichard to the Museum of Natural preserved at the Institut Français d’Afrique Noire History of Tripoli (Hufnagl, 1972) (Fig. 4). (CG. IFAN 44-24-5), and consisted in one skull e inscription on the label has faded but it is still with mandible, obtained from an unnamed site possible to read the date 1953, or perhaps 1958.

Fig. 4 - Present distribution of olive baboons and patas monkeys in north-, with the location of the mountainous massif of Tibesti, former homeland of Papio anubis tibestanus. Above left, the artistic illustration of a male baboon from the rock shelter of Tin Aboteka, located in south- western Tassili n’Ajjer, and stylistically referred to the archaic phase of the Round Head paintings. Above right, the skull of the Tibesti baboon on display in the Museum of Natural History of Tripoli, Libya (photo Marco Masseti). M. Masseti & E. Bruner 45

Groves (2005) places P. a. tibestianus among Ficus, and Myrtus, appears to be located around the synonyms of P. anubis. It is possible that the southern slopes of Tibesti up to the middle the Tibesti subspecies survived in some remote Holocene, albeit with some intrusions further wadies of the geographic area of the same name north (Sansoni, 1994). Proceeding hand-in-hand up to the end of the 1960s. In fact, up to 1969, with the process of aridifi cation, which revealed Beck & Huard (1969) still reported that: “Le a brusque acceleration in the late 4th millennium Teda, qui les appellent dongo ou dounbou, racon- BC, the tropical vegetation spread to the north, tent parfois qu’ils descendent d’hommes qui ont été progressively relegating the Mediterranean species ainsi transformés par Dieu pour avoir fait preuve to the higher parts of the mountainous massifs, de lâcheté” (=“ e Teda, who call them dongo where several specimens, regarded as multimillen- or dounbou, sometimes claim that they derived nial, have survived up to the present day (Sansoni, from humans and were thus transformed by God 1994). In fact, together with the endemic Ficus because they had shown cowardice”). According teloukat Battand & Trab., 1912 which grows on to Dorst & Dandelot (1973) and Halthenorth & the south and south-western slopes, the Saharan Diller (1977),baboons were still reputed as occur- mountainous vegetation of the upper reaches sup- ring in the in the 1970s. As a ports typical Mediterranean elements, such as general note however, lack of research in a region Nerium oleander (L., 1753) on the wetter north- does not mean necessarily that relic populations ern slopes (White, 1983). Archaeological evidence are now extinct. suggests a shift to the north of the borderline of the African savannah during the humid period of Note on the ecology of Tibesti the Holocene, probably in the 7th millennium BP Despite its geographical location in the (5th millennium BC) (Balout & Roubet, 1980). Saharan desert, the region of Tibesti is still char- Remnant tropical and Mediterranean spe- acterised by peculiar ecological conditions which, cies can be found throughout Tibesti, including up to few decades ago, consented the survival of palms, Hibiscus sp. and Rhynchosia sp. Others are the aforementioned baboon population. is was Saharan endemics with tropical or Mediterranean apparently a residual population from the period affi nities (White, 1983). ese species occur in in which this part of the Sahara had more suitable the area because the climate was wetter during the environmental and climatic conditions. In this Pleistocene and there was a continuous connection regard, it is interesting to note that the parallel of between this region, Mediterranean North Africa Tibesti (20°N) appears to play the role of biogeo- and . e past vegetation is docu- graphical threshold, since it also marks the north- mented by pollen grains found in the soils and ernmost limit of the natural dispersion of several sands of the desert and in the rock art of ancient plant species, such as the Higligh or “elephant people who depicted savannah woodland mam- tree”, Balanites aegiptiaca (L.), and the dom palm, mals, including African elephants, Loxodonta afri- Hyphaene thebaica (L.) (Petragnani, 1928). cana (Blumenbach, 1797), and various diff erent Formed of eroded volcanic rocks (Pritchard, species of bovids (Cloudsley- ompson, 1984). 1979), the pinnacled landform of Tibesti is still Several elements of this Sudanese fauna were able characterised by a residual vegetation of conifers to settle along the river valleys. e desert cat- and other as a remnant of the limited south- fi sh, Clarias lazera Cuvier & Valenciennes, 1840, wards expansion of the Mediterranean vegetation is an example of this Afrotropical fauna which in the course of the last glacial episode (Brown, extended its diff usion to the far North. Crocodiles, 1965). Since the ancient Holocene, however, the Crocodylus niloticus Laurenti, 1768, were also Mediterranean vegetation has been infl uenced by reported from the foothills of Tibesti (Duveyrier, the eff ect of arid degradation. e borderline with 1864; Lavauden, 1926; Brown, 1965). Among the the xerofi le vegetation of tropical type, such as sev- remnants of this sub-Saharan fauna we could also eral representatives of the genus Acacia, Tamarix, include the relic diff usion of baboons in Tibesti.

www.isita-org.com 46 The primates of the western Palaearctic

ere is, however, evidence for a former that currently exist within the distribution range occurrence of these primates in other Saharan of the Saharan cypresses, most signifi cant from areas too, such as the not distant mountainous a biogeographical point of view are the Saharan masses of Tadrart Acacus (Libya) and Tassili olive tree, Olealaperrini Batt. & Trab., 1912, and n’Ajjer (Algeria). the Saharan myrtle, Myrtus nivellei Batt. & Trab., 1912, both endemics of the mountainous com- The genus Papio in the archaeozoology of the plexes of the central Sahara and indubitably of Northern Sahara: Tassili n’Ajjer, Fezzan and Mediterranean origin (Charco, 1999). Tadrart Acacus Comprised, as noted above, within the politi- Primates are very rare among the and cal boundary of the Libyan Fezzan, the mountain- subfossil materials provided by the archaeological ous complex of Tadrart Acacus extends for 40 km investigation of the Sahara desert. We have already west to east, from the plateau of Tassili n’Ajjer at mentioned the few Neolithic bone remains of M. the western border of the Libyan Sahara, and for sylvanus yielded by some Algerian sites. Instead, 120 km north to south, from the southern slopes osteological fragments from Holocene deposits of Jebel Soda to the wells of El Uàr (Petragnani, in Dakhla Oasis (, Egypt) that were 1928). Located between 24°N and il 28°N, the initially recorded as “? monkey Cercopithecus” by Fezzan also falls completely within the boundary of Churcher et al. (1997) have since been identifi ed the Western Palaearctic, as indicated by Ellermann as Felis silvestris Schreber, 1775 (cf. Osborn & & Morrison-Scott (1951), and Corbet (1978). Up Osbornová, 1999). to 1930, Scortecci (1939, 1940) noted that typi- Scientifi c investigation provides evidence cal Afrotropical elements were unknown from the regarding the ancient Saharan dispersion of territories beyond the mountains of Sèrdeles, the baboons. It seems, in fact, that in former chro- northernmost part of Acacus, but were however nologies of the Holocene, these animals also found in the adjacent basin of Gat. Pollen analy- inhabited several territories located in the moun- ses carried out on several Holocene samples from tain ranges of the Tassili n’Ajjer (Algeria) and the Wadi Teshuinat (24°30’N 10°11’E), in the Tadrart Tadrart Acacus region in south-western Fezzan Acacus, on the whole, testifi ed a wetter climate and (Libya). Not unlike Tibesti, these areas too have a denser vegetation than today (Trevisan Grandi et been characterised by an indubitable richness of al., 1998). ey also documented a former time fl ora and fauna up to very recent times. Even span, from 6500 to 5500 BP, characterised by a in the fi rst half of the last century, according to greater availability of water, suffi cient to sustain Lavauden (1927 and 1930), in the inner part of a savannah-like vegetation, followed by a drier Tassili n’Ajjer, “dans la vallée de l’Oued Ihmirou et phase from about 5000 to about 3900 BP, when les vallées adjacentes il esiste des parties véritablement the process of aridifi cation began to be evident and boissées” (= in the valley of Oued Ihmirou and in the vegetation came to be confi ned in favourable other adjacent valleys…there are truly wooded water-receiving habitats such as wadies. Over the areas). A few dozen Saharan cypresses, Cupressus last century, these southern Libyan territories have dupreziana A. Camus, 1926, representatives provided extensive archaeological records, associ- of Mediterranean vegetation in the mountains ated with the human population spanning from of central Sahara, still survive today in the sur- the Mesolithic (9000-7000 bp) to historical eras roundings of Tamrit (Borzatti von Loewenstern, (Di Lernia & Manzi, 2002). In particular, this area 1982), and, albeit with a much more limited preserves many cave and rock paintings and draw- presence, on the mountainous massif of Ahaggar. ings, enhancing the available information on the ese trees, some of them millennial, are now lifestyle of these human populations, their social regarded as authentic living that survive as structures and processes, their relationship with best they can in the extreme climatic conditions the territory and fauna (Mori, 1965). Although of the region (Charco, 1999). Among the species these territories of southern Libya have furnished M. Masseti & E. Bruner 47

numerous osteoarcheological records, monkey majority of the fi nds emerging from the archaeo- bones are nevertheless very scarce. Some remains logical exploration of this area, it may be attribut- were originally described as Cercopithecus aethiops able to the seventh-sixth millennium BP. (Cassoli & Durante, 1974) at Uadi Ti-n- ora, and More recently, a fragment of maxillary bone with dated at around 8000-8500 years bp (Gautier & a third molar was recovered at the site of Takarkori, Van Neer, 1977), but they have recently been clas- again by the Italian-Libyan Archaeological Mission sifi ed as P. cynocephalus (L., 1766) (Gautier & Van from the “La Sapienza” University of Rome. In line Neer, 1977). Ordinarily referred to as the yellow with the general morphology, and above all the baboon, this latter species is slimmer than P. anubis, main diameters reported for the African primates and is now widespread in sub-Saharan Africa, from (Swindler, 2005), again it can only be assigned to the river Zambesi to Tanzania, coastal Kenya and the genus Papio (Bruner, unpublished data). Here, (cf. Hill. 1966; Kingdon, 2004; Groves, several burials have been identifi ed in a rock shelter, 2005). us, in its northern and easternmost diff u- and the specimen can be dated to the Mesolithic sion, the extends through the terri- (9000-7000 ybp), or the early Pastoral (7000- tories of East Africa but is completely missing from 6000). e former phase refers to hunter-gathering the biogeography of the Sahara desert proper; as far and fi shing, with an economy largely based on the as is known at present. It also appears to be practi- Barbary sheep Ammotragus lervia (Pallas, 1777). cally unknown in Saharan palaeozoogeography. Up e latter phase is associated more with domes- to a few years ago, however, various populations of tication and domestic caprine resources. Taking baboons were still comprised within the scientifi c into account the dimensions and biogeographi- defi nition of this species. Only recently have several cal ranges of the currently recognised baboon of these instead been assigned to diff erent taxa (cf. taxa (Groves, 2001), both the Takarkori and Uan Groves, 2001 and 2005). Considering, in fact, the Kasa specimens can be provisionally assigned to P. extant range of the distribution of the North African anubis. e same identifi cation should be applied representatives of the genus Papio, the taxonomic to the specimens formerly assigned to P. cynocepha- assessments clearly refer to the systematic single- lus in the same areas or in similar archaeological species interpretation of the genus Papio, now split contexts. Even if the presence of baboons in these into at least fi ve diff erent taxa (cf. Groves, 2001). P. areas is further confi rmed, it nevertheless appears cynocephalus was also described by Gautier (1987b) that there was no relevant relationship with the in the fauna from Uan Muhuggiag in the Pasa col- human settlements. It remains to be investigated lection at the Museum of Verona. A few osteologi- whether or not these primates were associated with cal and dental fragments were further reported dur- human activities (hunting, pets), and whether they ing an archaeological survey on superfi cial layers at were introduced from other regions or represented the site of Uan-Kasa (sample TH 131), and again residuals of the prehistoric and/or protohistoric described as P. cynocephalus by Corridi (1998). baboon distribution. According to Claudio Corridi (pers. com., 2007), the sample included one canine and two other Representation of primates in prehistoric Saharan art undetermined osteological fragments. e fi nd Indirect evidence of the former occurrence came from the site of Sennadar (24°54’N 10°28’E), of primates - and in particular of baboons - in where it was found in a shelter located in the wadi Saharan zoogeography is also furnished by artis- alluvium in the course of a surface survey in Acacus tic productions, although such representations and Messak performed between 1990 and 1993 are few (cf. Gautier, 1993). In reality, monkeys by the Italian-Libyan Archaeological Mission from rarely fi gure in the iconographic records of local the “La Sapienza” University of Roma. is mate- rock art. Carved representations of monkeys at rial was supposed to be related to a chronological Hadjrat Driess (Hachid 1992), and Djorf Meharra phase comprising between the Mesolithic period (Soleilhavoup, 2003) have been recorded from the (Late Acacus) and the Middle Pastoral, but like the Atlas mountains.

www.isita-org.com 48 The primates of the western Palaearctic

e artistic illustration of a primate is also shelter of Oued Tirehart, in western Tassili (Kunz, recorded from the rock shelter of Tin Aboteka, 1988). e ethnozoological interpretations are not located in south-western Tassili n’Ajjer, along always univocal (Plate 8). At In Farden there is an the Algerian slope of Oued Edjériou (see Fig. 4). alleged representation of a red monkey which is e image consists of a large profi le painted in hunting, but the image is very stylised, and diffi - a white colour with a red outline. It could eas- cult to assess (Mori, 1965). is representation has ily be identifi ed as the representation of an adult been connected with the artistic context of the so- male baboon, portrayed with an aggressive atti- called Late Pastoral style. Close to this site, among tude (cf.Napier & Napier, 1967; Despard Estes, the artistic production of the Round Head style, 1991). Stylistically it has been included in the another monkey profi le was initially recognised archaic phase of the so-called Round Head paint- at Ti-n-ascig, but the image is more likely to be a ings (Sansoni, 1994). is production is stylis- large carnivore (Mori, 1960 and 1965). Carnivores tically well-defi ned, although its chronological and baboons are easily misinterpreted in rock art location remains somewhat vague. e bulk of because of the many similarities associated with the the production belongs to the period comprised body profi le: enlarged chest, certain positions of within the “Neolithic Wet Phase” (c. 4500-2500 the tails, the muzzle and the canines, and for some BC) and before the “Postneolithic Arid Phase” species also the mane. Nevertheless, it appears that (c. 2500-1000 BC) (Muzzolini, 1989). us, a group of four baboons can be easily recognised in Round Head paintings can be chronologically a wall drawing at the site of Teshuinat III (Mori, located between the unspecifi ed commencement 1965), possibly dating to the Middle Pastoral of human settlement of the last Saharan humid (6000-5000 ybp). e mane and sexual character- phase (dating perhaps to the 8th millennium BC) istics suggest that the fi rst is an adult male, followed and the arid pulsation of the 6th millennium BC by three juveniles or females (Plate 8). Mori (1965) which, according to Sansoni (1994), anticipates interpreted the images as hamadryas, remarking the advent of the food production economy on that although these baboons live in sub-desert a large scale. It is also interesting to note the ele- rocky habitats their presence in Acacus rock art is vated affi nity between the zoological species rep- fairly rare. In any case, in terms of biogeography resented in the Round Head artistic productions they can most probably be interpreted as P. anubis. (Kunz, 1988; Muzzolini, 1989; Sansoni, 1994) More generally, one can observe that, among the and the osteological material provided by the few certain images of monkeys in Saharan art, it archaeological exploration of the contemporary appears that the only species portrayed was the sites of the Libyan desert, in the Fezzan-Acacus olive baboon. Muzzolini (1989) is of the opinion (Cassoli & Durante, 1974; Gautier & Van Neer, that in Round Head art these monkeys are to be 1977; Gautier, 1987a, 1987b and 1993; van regarded as “symbolic” animals. der Veen et al., 1996; Corridi, 1992 and 1998; Alhaique, 2002). Characterised by a very old patina, and included in the earliest sub-group of or sacred baboon, the same artistic production, is the illustration Papio hamadryas (L., 1758) of what is probably a baboon from the site of Tizzeine in Central Tassili (Muzzolini, 1989). French: hamadryas Among the artistic productions of both Tassili Spanish: hamadríade n’Ajjer and Tadrart Acacus, few other represen- Italian: amadriade tations of monkey related to the Round Head German: Mantelpavian group are known. ere are other baboons - or pseudo-baboons, Olive baboons are absent from north-eastern for some of them have human limbs, or even Africa (Ogaden, northern Somalia) (Funaioli, appear to be holding a bow – in the paintings of the 1971; Haltenorth & Diller, 1977; Yalden et al., M. Masseti & E. Bruner 49

1996). ese territories are, instead, the home- The sacred baboon of ancient Egypt land of another representative of the genus Papio, Smith (1969) estimated that the sacred the Hamadryas baboon or sacred baboon, Papio baboon became extinct in Egypt by the third hamadryas (l., 1758), dispersed in the arid zone millennium BC or the First Dynasty. According of the Red Sea coast of Sudan, Eritrea, to Arnold (1995), it vanished from the wild and northern Somalia (Hill, 1970; Funaioli, during the Middle Kingdom (2134-1785 BC): 1971; Haltenorth & Diller, 1977; Al-Safadi, thereafter baboons continued to be imported 1994; Yalden et al., 1996; Groves, 2005). is from the south. Kummer et al. (1981) are of the species occurs in two populations which are now opinion that it is quite likely that the Egyptians completely separated by the Red Sea. In fact, it imported these animals from African latitudes is found both in Africa and in Arabia, where it further south than their own country. In any lives in the mountainous south-western corner of case, no fossil remains of the species have been the peninsula, occurring on the Red Sea hills in found in Egypt and there is no defi nite knowl- south-western Saudi Arabia and western Yemen, edge of its ancient distribution. e baboon cult, particularly near Aden ( omas, 1900; Elliot however, apparently began in the Predynastic 1913; Starck and Frick 1958; Harrison, 1964; period when it seems that this primate existed Kummer et al., 1981; Nader, 1990; Harrison & in Egypt (Osborn & Osbornová, 1999). It is Bates, 1991; Al-Jumaily, 1998). is was also one believed that hamadryas were generally kept in of the species of monkey best known in antiquity. temples and embalmed. ey rarely became pets, It has at length been believed that the diff usion because of their aggressive nature (cf. Erman, of hamadryas in Arabia may have been related 1894; McDermott, 1938). Representation of to their importation from the opposite coast of sacred baboons is found in artistic produc- Africa from at least the time of ancient Egypt (see tion from the Protodynastic chronologies (end Kummer et al., 1981). is is the only species of of the 4th millennium-2695/2640) up to the baboon taken into consideration by Ellermann Christian era, in which period the worship of the & Morrison-Scott (1951), but, as the authors baboon persisted (Osborn & Osbornová, 1999). explain, their work on the Palaearctic and Indian Particularly famous are the hamadryas evoked mammals is limited to the African territories in the wall-paintings of Tutankhamun’s burial located north of the 20° parallel, although they chamber, where they are supposed to guard the deal with the whole of the Arabian peninsula. young pharaoh as he passes through the 12 sec- Corbet (1978) also regards the hamadryas as the tions of the underworld on the fi rst night after only baboon occurring in the Palaearctic region. his death (Plate 9). We are dealing with a species whose natural It appears that the major period of primate distribution now appears to be limited to the importation was the New Kingdom, which north by the 20°S parallel, which – as mentioned began with the XVIII Dynasty (second half of above - is regarded as the southernmost limit of the 2nd millennium BC) (Osborn & Osbornová, the Western Palaearctic unit. It has, however, been 1999). In several artistic productions of this suggested that in antiquity hamadryas baboons period, it is possible to detect the southern origin were dispersed further north, up to the territories of the imported monkeys, which were normally of Sudan, and even Egypt. e latter coun- brought from Nubia (i.e., the wall-paintings of try is also regarded as the “type locality” of the the tomb of Rekhmire at ebes) and the land species (Napier, 1981; Groves, 2001 and 2005), of Punt (i.e., the reliefs of the tomb of Queen even though sacred baboons have long since van- Hatshepsut at Deir el-Bahari). It is not known ished from these territories. In any case, Linnaeus whether Punt lay on the African or the Arabian described the taxon in his Systema Naturae (1758) coast. Its location is supposed to be along the through the examination of specimens from two shores of the Straits of Bab el Mandeb, at the “Egypt” and “” (cf. Groves, 2001). end of the Red Sea, and along the coast of north-

www.isita-org.com 50 The primates of the western Palaearctic

eastern Somalia and southern Arabia (Osborn & is, however, the only baboon whose range extends Osbornová, 1999). Punt, according to Erman beyond continental Africa (cf. Napier & Napier, (1894), “evidently signified the more tropical coast 1967). According to Haltenorth & Diller (1977), lands of the Red Sea, the south of Arabia, and the the hamadryas is dispersed in the mountainous Somali coast”. areas along the coasts of the south-western and According to Osborn & Osbornová (1999), southern regions of Arabia and Yemen, about 20° information regarding primate mummies is N to about 50°E (Nader, 1990; Al-Jumaily, 1998). somewhat confusing, mainly because of the lack In the peripheral hills of these areas, the baboon of supporting data. Various authors, including has recently become abundant (Corbet, 1968; MacDonald (1965), and Rabb (1968), stated for Harrison& Bates, 1991). In fact, its distribution example that thousands of hamadryas had been is strongly infl uenced by the availability of food, mummifi ed because they were so highly vener- water, and safe havens (Al-Safadi, 1994). us, it ated and respected. Others, such as Brunner is the only large that can be considered (1969), remarked on the fi ndings of thousands to be over-abundant, causing problems to farmers of mummifi ed ibises at Saqqara that brought and local people (Abuzinada et al., 2002). e rel- the species close to and “similar fates ative abundance of baboons is not aff ected by the befell the cats in Bubasti …and baboons in Tuna el presence of predators. However, no troops have Gebel ”. Boessneck (1988), however, noted that been recorded from northern Yemen (Al-Safadi, the baboon mummies were in any case much fewer 1994). ere is also evidence for the occurrence than the ibis mummies, for in the Late Period no of sacred baboons on some of the islands of the more baboons of either species were imported and Red Sea, such as the Farasan archipelago. A com- they were no longer kept in temples. plete skull with mandible of a subadult female of the species was in fact discovered in a provisional The Arabian sacred baboon, Papio hamadryas burial on Farasan Al-Kebir, the largest island of arabicus (Thomas, 1900) the archipelago (Fig. 5). is island is located Arabian baboons have been provisionally opposite the south-western coast of Saudi Arabia, ascribed to Papio hamadryas arabicus ( omas, some 50 km off shore from the town of Jizan, at 1900) (Ellermann & Morrison-Scott, 1951; 16°42’21’’N 41°59’0’’E. e sacred baboon skull Harrison, 1964; Corbet, 1978; Harrison & Bates, was collected in April 1984 by the Italian zoologist 1991). is subspecies was created by omas Benedetto Lanza, in the course of the joint mis- (1900) on the basis of the examination of an adult sion of the Gruppo Ricerche Scientifi ce e Tecniche female specimen collected, on 16 October 1899, Subacquee of Florence and the Feal Costruzioni in Subaihi Country, about 60 miles north-west of of Milan, which was building a hospital there at Aden at an altitude of 1000 metres a.s.l.. Its type the time. e specimen is now preserved in the specimen is still conserved in the collection of the collections of the Museum of Natural History of Natural History Museum of London, (BMNH the University of Florence, Zoological Section 99.11.6.1.) According to Harrison (1964), the “La Specola”, under catalogue number MZUF material available is, however, hardly adequate to 11329. A transfer of pets by humans is the most assess the validity of the Arabian race, although it likely explanation of this occurrence of a hama- is possible that it may perhaps be distinguishable dryas baboon on Farasan Al-Kebir. by an, on average, smaller size and smaller cheek- Regarding the present disjunction of the range teeth. In a preliminary study of the behaviour and of the hamadryas, the Red Sea acts as an extrinsic ecology of four Arabian sacred baboon popula- barrier to gene fl ow between continental and pen- tions in south-western Saudi Arabia, Kummer et insular hamadryas populations (Wildman, 1999). al. (1981) also concluded that the animal should But Kummer et al. (1981) observed that there not be considered a separate subspecies from the is at present no evidence of a genetic diff erence north-east African one. e Arabian hamadryas between Arabian and Afrotropical hamadryas M. Masseti & E. Bruner 51

baboons. ere is no north-south gradient of African galliform is restricted to the south-west- morphology or behaviour in Saudi Arabia that ern portion of Arabia (Silsby, 1980; Gasperetti, could give a clue as to where the two populations 1981), but the lesser kudu was an African ungu- had their most recent contact. e most recent late unknown in the until Harrison land bridge across the straits of Bab el Mandeb, at (1972) and Büttiker (1982) recorded two separate the southern end of the Red Sea, existed around specimens: the fi rst from Yemen and the second 12-10,000 years ago (cf. Bailey et al., 2007), and from the Medina province (Saudi Arabia). is even then it may have been a salt desert diffi cult ungulate inhabits the arid thornbush country for baboons to cross. Moreover, the absence of P. areas of East Africa from Ethiopia to Tanzania, up hamadryas from Oman suggests an African ori- to 1300 m (Funaioli, 1971; Haltenorth & Diller, gin of the species. Looking for a more recent pos- 1977), and, as of today there is no palaeonto- sible exchange, there is a theory that hamadryas logical evidence of its ancient presence in south- baboons were imported by the ancient Egyptians, western Asia. us, as in the case of the - to whom they were sacred, from the . fowl, it could be possible that the lesser kudu was As already mentioned, it is still unclear whether previously introduced from East Africa to Arabia Punt lay on the African or the Arabian coast of the as a game animal (Borzatti von Löwenstern & southern Red Sea, but in either case it is probable Masseti, 1991). Kummer et al. (1981) is of the the sacred baboon was transferred from one coast opinion that ancient vessels, or even a more recent to the other (Kummer et al., 1981). Over recent transfer of pets by humans, might still be the most decades, other biological elements of Afrotropical likely explanation of the occurrence of hama- origin, such as the guinea fowl, Numida meleagris dryas on both Red Sea coasts. However, recent (L., 1758) and the lesser kudu, Tragelaphus imber- genetic analyses, performed by Lawson Handley bis Blyth, 1869, have also been reported from et al. (2006), confi rmed no signifi cant diff erence Arabian zoogeography. e occurrence of the between the African and Arabian populations

Fig. 5 - Sacred baboons occur in two populations which are completely separated by the Red Sea, being found both in Africa and in Arabia. There is also evidence for the occurrence of this species on the Farasan archipelago, where a complete skull was discovered on Farasan Al-Kebir, the largest island of the archipelago (16°42’21’’N 41°59’0’’E) (photo Saulo Bambi; courtesy Museum of Natural History of the University of Florence, Zoological Section “La Specola”).

www.isita-org.com 52 The primates of the western Palaearctic

examined during the study. Consequently, it is 1990). Since only a single species of Erythrocebus unlikely that the low level of variation on the Y is now recognised, the geographical range is the chromosome found in Arabia was caused by a same of that of the genus (Hill, 1966). eir population bottleneck during the colonisation of distribution ranges from the Sudanese zone of Arabia by African hamadryas baboons. Senegal and Mauritania to the Upper and the Some years ago, Wildman (1999) used Atbara in the east (northern border between 18° mitochondrial DNA sequences and nuclear and 15°N, southern limit in the west and centre short tandem repeat genotypes to estimate the around 10°N, in the east around 3°S; Haltenorth dates of dispersal of African hamadryas popu- & Diller, 1977; Groves, 2005) (Fig. 6). Hill lations onto the Arabian peninsula. e results (1966) recognised at least four subspecies of this of this study included sequences and genotypes taxon: Erytrocebus patas patas (Schreber, 1775), of hamadryas baboon populations from Yemen, E. patas villiersi (Dekeyser, 1950), E. patas pyr- Saudi Arabia, and Ethiopia, as well as data from ronotous Hemprich & Ehrenberg, 1829, and E. appropriate papionin outgroup taxa. Genetic patas baumstarki Matschie, 1905, the latter form data were analysed using philogenetic and being confi ned to a restricted area on the south- population genetic techniques, and the analy- eastern shore of Lake Victoria. However, accord- ses suggested that a dispersal onto the peninsula ing to Groves (2001), the taxonomy of this spe- occurred during the Middle Pleistocene. When cies is still imperfectly known: subspecies may combined with geological, climatological, bio- exist, but at least some of the features that alleg- geographic and historical evidence, the results edly characterised them were based on changes raise several points of discussion. Hamadryas in the female’s facial patterns during pregnancy. baboons were not introduced onto the Arabian e subspecies E. patas patas Schreber, 1775, peninsula by Egyptians or other historic human commonly called the West African red monkey, groups. e animals may have instead dispersed ranges for example to the north-east onto the across a now submerged isthmus at what today Ennedi plateau (north-eastern Chad), and as far are the straits of the Bab al Mandeb at the north as the Mourdi depression. It has also been southern end of the Red Sea, rather than across reported in the northern part of Ubangi (Hill, the Sinai peninsula. us, it seems that humans 1966; Napier, 1981). Patas monkeys are appar- were not the only primates to migrate out of ently capable of surviving for long periods with- Africa during the Middle Pleistocene (cf. Rook out water. Mason (1936), for example, shot one et al., 2004). specimen north of Wadi Hawar, in north-western Sudan, and also found skulls of the species in the area “…hundreds of miles from any water”. or red monkey, According to Osborn & Osbornová (1998), Erythrocebus patas (Schreber, 1775) patas monkeys probably existed in Egypt, in what is now desert, into the early dynasties (end French: patas, singe rouge of the 4th-3rd millennium BC). ey were per- Spanish: patas, mono rojo haps popular as pets during the dynastic period, Italian: eritrocebo, scimmia rossa but their alleged artistic reproductions are diffi - German: Husarenaffe cult – or almost impossible - to distinguish from those of other long-tailed monkeys. e archaeo- ese are highly distinctive medium-sized zoological evidence is also scarce. Kessler (1989), monkeys, with long limbs and a slender build for example, listed only one patas monkey from (cf. Haltenorth & Diller, 1977; Kingdon, 1994). the Late Period (beginning of the 2nd-1st mil- Erythrocebus patas (Schreber, 1775) is a Sahelo- lennium BC) animal necropolis in Dendera. e Sudanese species, some insularised populations of Greek poet Aelian (c. 170- after 230 AD) left which subsist in the Sahara (Hill, 1966; Le Berre, us a fascinating and accurate description of the M. Masseti & E. Bruner 53

external morphology of this animal: “In his writ- 1992), which we feel must be included among ing about the Red Sea Pythagoras says that there the primates of the western Palaearctic. However, is an animal that lives on the shores and is called as noted above, we should not forget that Vaurie Kêpos. And it is well-named (kêpos, garden), for (1959-1965) and Cramp (1977) include the it is of many colours. When full-grown it is the size mountain massif of Tibesti include within the of an Eritrean hound. But I wish to return to the confi nes of the Western Palaearctic, but exclude subject of its varied colouring and to describe it as those of Aïr (Niger) and Ennedi (Chad), due to he writes. Its head, its back, and its spine down as the predominance of the Afrotropical biogeo- far as the tail are pure red, though you may observe graphical element in the latter territories. e a sprinkling of golden hairs. But its face including Aïr population of patas monkeys was described the cheeks is white, and from there golden stripes for the fi rst time by Dekeyser (1950) who indi- descend as far as the neck. The lower portions down cated its members as belonging taxonomically to its chest and its forefeet are all white; its two to the new subspecies Erythrocebus patas villiersi breasts, which would fill your hand, are dark, but (Dekeyser, 1950), commonly referred to as the its belly is entirely white; its hind feet are black ” Aïr patas monkey (Type Specimen: CG IFAN (On the characteristics of animals, XVIII: 8 [cf. 47-10-165, ♂adult) e type locality of this taxon Scholfi eld, 1959]). is Irabellaben, where it was reported from an alti- tude of 1,200-1,300 m. Regarded as native to Aïr, Aïr patas monkey or Aïr red monkey, the taxon is characterised, inter alia, by reduced Erythrocebus patas villiersi (Dekeyser, 1950) dimensions compared to the nominal form and Patas monkeys have also been recorded from by longer canines (Dekeyser (1950 and 1955; between 15°-20° N (Napier, 1981). e semi- Napier, 1981). Nevertheless Hill (1966) consid- Sahelian biocenoses of the Aïr massif region ered the exceptionality of these long canines as a (Niger), in fact, still permits the survival of an possible individual anomaly. e Aïr population isolated population of these monkeys, (Bousquet, is isolated from the Sahelian range of the species

Fig. 6 - The distribution of patas monkeys ranges from the Sudanese zone of Senegal and Mauritania to the Upper Nile and the Atbara in the east; northern border between 18° and 15°N, southern limit in the west and centre around 10°N., in the east around 3°S. (from Hill, 1966).

www.isita-org.com 54 The primates of the western Palaearctic

and is itself composed of several discontinu- updated, moving all the species from the genus ous and isolated pockets (Dekeyser & Derivot, Cercopithecus to a new genus, Chlorocebus (Rowe, 1959). Dekeyser (1950) states that this monkey is 1996; Groves, 2001, 2005). ere are now at common practically throughout the region of Aïr, least six representatives of this genus recognized inhabiting the larger valleys of the south. Its alti- at species level, with a geographical range extend- tudinal range is between 600 m. a.s.l. at Dahaga ing over most of sub-Saharan Africa from about and 1,600 m. on the plateau of Baguezans. 18°-15°N, although excluding a large part of the south-western, southern and south-eastern conti- Nisnas, Nile patas, dancing red monkey or nental landmass (cf. Haltenorth & Diller, 1977). Blue Nile hussar monkey, Erythrocebus patas In mainland , the northernmost limit pyrrhonotus Hemprich & Ehrenberg, 1829 of these monkeys was given by Rode (1938) as St. Another population of patas monkey has Louis at the mouth of the Senegal river. However, been reported from Wadi Hawar, in north- Galat & Galat-Luong (1977) recorded a popula- ern (north-western Sudan). Comprised tion occurring slightly further north, on the island between 15°N and 20°N, this basin lies near the of Morfi l (16°33’N 14°44, 5 E), located in the border with Libya and Chad, between the junc- valley of the river Senegal, south of the village of tion of the east-west route linking the Sahara Podor, near the border with Mauritania. Groves and the Nile Valley and the north-south route (2005) recognises the following species: the grivet linking northern Africa and the Sahel-savannah monkey, Chlorocebus aethiops (L., 1758), distrib- belt (cf. Mohammed-Ali, 1981). As noted above, uted in Sudan east of the White Nile, Eritrea, nisnas were found by Mason (1936) about 300 Ethiopia east of the Rift Valley; the , miles from the nearest open water. Hemprich C. cynosurus (Scopoli, 1786), which spreads from & Ehrenberg (1829) ascribed it to the taxon E. the southern Democratic Republic of Congo to pyrrhonotus (cf. Napier, 1981), but today it is northern , and west of the river regarded as an authentic full subspecies, E. p. Luangwua; the Bale Mountains vervet, C. djam- pyrrhonotus Hemprich & Ehrenberg, 1829, dis- djmanensis (Neumann, 1902), diff used over the persed from Sudan, Atbara and western Ethiopia highlands east of the lakes Abiata, Shalla and to western Kenya and (Mason, 1936; Zway, in Ethiopia; the , C. pyger- Hill, 1966; Haltenorth & Diller, 1977; La Berre, hytrus (F. Cuvier, 1821), dispersed in Ethiopia east 1990; Groves, 2005). e primate was described of the Rift Valley, Somalia to Zambia east of the as having a completely white nose spot. Later, river Luangwua, and ; and the tan- Pocock (1907) realised that the colour of the talus monkey, C. tantalus (Ogilby, 1842), whose nose-spot varied with age and Loy (1974) men- distribution ranges from the river Volta (Ghana) tioned the eff ect of the animal’s reproductive east to the White Nile (Sudan) and Lake Turkana condition upon this. (Kenya). Finally, the green monkey or Cape Verde monkey, C. sabaeus (L., 1766), is naturally distrib- uted from Senegal to the river Volta (Ghana), also Green monkey or Cape Verde monkey, occurring on the islands of Cape Verde (Haltenorth Chlorocebus sabaeus (L., 1766) & Diller, 1977; Groves, 2005) (Fig. 7). e lat- ter is an archipelago approximately 500 km off French: singe vert, singe vert du Sénégal the west coast of Africa at 15°02’N, 23°34’W, Spanish: tota formed by ten main islands and about 8 islets Italian: cercopiteco gialloverde which are divided into two groups: Barlavento, German: Gelb Grünmeerkatze the northern island group, and Sotavento, the southern island group. Due to its location in the e taxonomic classifi cation of guenon eastern Atlantic Ocean, this archipelago was not monkeys, or vervet monkeys, has recently been included by Ellermann & Morrison-Scott (1951) M. Masseti & E. Bruner 55

and Corbet (1978) within the boundary of the C. sabaeus was taxonomically separated from Western Palaearctic. But, as mentioned above, , C. aethiops (L., 1758), as a full species Vaurie (1959-1965), de Naurois (1969 and by Kingdon (1997) and Groves (2001). Groves 1994), and Cramp (1977) suggest comprising it (2005) indicates the archipelago of Cape Verde as within the confi nes of this zoogeographical region its type locality (terra typica). In fact, the descrip- in view of the evident Palaearctic character of its tion of Linnaeus and the Latin name of the spe- avifauna. Several other authors, such as Coutinho cies, sabaea, (Systema Naturae, 12th ed., 1: Saraviva (1961), Sunding (1970 and 1979), 38) were based mainly on a single specimen orig- Kunkel (1980), Ribeiro et al. (1980), Gonzales inating from the island of Santiago, which was Henriquez et al. (1986), and Beyhl et al. (1995) published by George Edwards in his Gleanings have also expressed themselves in favour of the of Natural history (1758-1764) (Osman Hill, inclusion of these islands within the limits of the 1966). Edwards (1758-1764) labelled his animal Western Palaearctic. the “St. Jago Monkey” since it was brought to

Fig. 7 - The continental distribution of guenon monkeys, or vervet monkeys, ranges from the terri- tories south of the Sahara and the Sahel to . Green monkeys or Cape Verde monkeys, Chlorocebus sabaeus (L., 1766) (above left - photo Claudio Vergano and Paola Lovesio) were also introduced in historical times on the archipelago of Cape Verde (below left) where they are today present only on the island of Santiago and, perhaps, Fogo (grey arrows: current distribution; empty arrows: historical distribution).

www.isita-org.com 56 The primates of the western Palaearctic

England alive from one of the Cape Verde islands & Zava (1988) are instead of the opinion that of that name (St. Jago) (Plate 10). e same this importation occurred from Senegal towards author said that the sailors of his time generally the end of the 19th century. Naurois (1994) referred to the green monkey as the “St. Jago and Sorgial (1995) observed, however, that the monkey” because the animals were brought to monkeys were deliberately released in Santiago, England from St. Jago in Cape Verde. Later on, while their release in Furna (Brava) was involun- both Jardine (1833) and Goldsmith (1840) noted tary, the result of an accidental escape on the part that green monkeys lived on the islands of Cape of the animals. In any case, the monkeys cannot Verde. Ogilby (1838) confi rms that the species have been released on Cape Verde before 1462, to which the name came to be applied was that the year of the discovery of the Atlantic archi- otherwise known as the green monkey and Cape pelago by the Portuguese (de Vasconcelos, 1920; Verde monkey. In the course of the 20th century, De Oliveira Boléo, 1939). these primates were still reported from the Cape Today there are plenty of green monkeys in Verde islands of Santo Antão and Brava (Muzio, Santiago (Hazevoet, 1995; Payne, 2003), while 1925), Brava and Santiago (Naurois, 1994; those introduced into Brava have not survived Hazevoet, 1995; Sorgial, 1995). More specifi - (Hazevoet, 1995). A large colony of the pri- cally, Muzio (1925) described these insular mon- mates lives in the palm grove behind the beach keys as “…a variety of little monkey (cercopithecus of Tarrafal, in the northernmost part of Santiago saboeus) to be found jumping around in the for- (Righetti, 2004). Green monkeys on Fogo are ests of Antao and Brava”. According to Naurois now kept as pets. Apart from Santo Antão, the (1994), the sub-species occurring in the archi- former and present occurrence of green mon- pelago is Cercopithecus aethiops sabaeus (L., 1766) keys seems to be restricted to the islands of the [= Chlorocebus sabaeus (L., 1766)]. Together with southern group of the archipelago, the Sotavento that of other exotic animals such as rats, sheep, group, possibly due to the more favourable cli- goats and cattle, the introduction of this monkey matic and environmental conditions, together too is considered to have had negative eff ects on with the presence of some wooded areas. the native fl ora and fauna of the archipelago. It seems very likely that the green monkeys African green monkeys in the of Cape Verde were introduced there from the (Neotropical biogeographical region) adjacent African mainland (Hill, 1966). In fact, In the late seventeenth century, green mon- it should be stressed that there is no trace of the keys were introduced into several West Indian osteological remains of these animals in Cape islands, in the western Atlantic Ocean, when Verde Pleistocene deposits. Nor does it seem likely ships involved in the slave trade travelled to that C. sabeus reached the Atlantic archipelago by the from West Africa (van der Kuyl swimming, jumping onto fl oating logs or other et al., 1996). Today, large feral populations of so-called sweepstake routes. It is thought that the these primates live in Barbados, St. Kitts, and primates were imported from continental Africa Nevis (Ashton & Zuckerman, 1950; Hill, 1966; onto the Cape Verde archipelago no later than Haltenorth & Diller, 1977), and small numbers the mid-17th century (Napier & Napier, 1967), have been reported from Sint Eustatius (Denham and perhaps even much earlier (Denham, 1987). (1987) (Fig. 8). e islands of St. Kitts, Nevis In fact, in 1673 during a visit to the already and Sint Eustatius are clustered together near the mentioned island of St. Jago, Fryer (1909) met northern end of the Lesser Antilles; Barbados is natives on the beach selling monkeys, “such green instead located about 600 km to the southeast of ones as are commonly seen in England to be sold”. that cluster, about 150 km from Grenada and the According to Naurois (1994) and Sorgial (1995), Antillean Volcanic Arc, and about 4000 km west the monkeys were imported respectively from of the nearest point of Africa (cf. Denham, 1987). Guinea and Guinea Bissau. Azzaroli Puccetti In some instances these monkeys have escaped M. Masseti & E. Bruner 57

and are now living as wild populations on some (1987) suggests that these West Indian mon- of these islands, notably St. Kitts, one of the keys are of heterogeneous origin, coming not Leeward Islands (Ashton & Zuckerman, 1950), only from Cape Verde but also from continental and Barbados (Hill, 1966). Specimens from St. West Africa. eir story is somehow connected Kitts are conserved in the collection of the Natural with that of the early European settlements of St. History Museum, London (BMNH), and the Kitts, Nevis and Barbados, the African origin of Odontological Museum of the Royal College West Indian slaves, and the transatlantic routes of Surgeons of London (Ashton, 1959; Ashton from the seventeenth century up to the nine- & Zuckerman, 1950; Ashton & Zuckerman, teenth, and even earlier. We also have to consider 1951a; Ashton & Zuckerman, 951b; Ashton the role of Barbados as a slave entrepot, the fact & Zuckerman, 1951c; Colyer, 1948a; Colyer, that monkeys were embarked as pets aboard ships 1948b; Napier, 1981; Sclater, 1866). and as trade goods sold to menageries in Europe Caribbean-born African green monkeys were and the New Word. In this respect, it is perti- classifi ed as C. sabaeus also through cytochrome nent to recall that there is clear evidence for the b sequencing (Pandrea et al., 2006). Denham importation of other mammals to the

Fig. 8 - Location of the West Indian islands, in the western Atlantic Ocean, inhabited by African green monkeys (black arrows), and the Cape Verde archipelago, off the coast of western Africa. Following the trade of slaves from Africa, another species of African monkey reached these Neotropical islands in recent historical times: the , Cercopithecus mona (Schreber, 1774) (white arrows). A population of mona monkey is reported from the Caribbean island of Grenada, in the Windward archipelago (Lesser Antilles) (Denham, 1987), where the species was probably introduced in the course of the eighteenth century (Napier, 1981). According to Haltenorth & Diller (1977), this spe- cies was also imported onto St. Kitts. In recent historical times, mona monkeys have also been introduced onto the Atlantic islands of São Tomé and Principe, in the (Frade, 1956 ; Afonso-Roque & Santinho Barata, 1992).

www.isita-org.com 58 The primates of the western Palaearctic

West Indies, even of larger size. In fact, in 1857, specimen must have been part of the menagerie Charles Darwin presented the British Museum of Pope Leo X who, like his late father Lorenzo (Natural History) with three skins of European de’ Medici, collected rare and exotic animals fallow deer, Dama dama dama (L., 1758), originating from all corners of the world (Bellori, from the island of Bartudee (Barbuda), near 1931; Fontoura da Costa, 1937; Dacos, 1969, Antigua, providing the following information: 1977; Clarke, 1986; cf. Masseti, 1991). A speci- “Domesticated and naturalised about 100-150 men of the same species of monkey fi gured later, years ago on the Island of Barbuda” (Chapamn & in 1794, again in Florence, among the collections Chapman, 1980; Masseti, 1996). ese skins are of the R. Museo di Fisica e Storia Naturale, from still preserved in the Natural History Museum of where it was transferred from the menagerie of London (BMNH 57.1.14.1, BMNH 57.1.14.2, the nearby Boboli gardens. It was listed under BMNH 57.1.14.3). e archipelago of Cape catalogue number 1129, with the name of Simia Verde was, indeed, a vital link in the British sabea (Sabean monkey) or Scimmiotto verde (= “triangle trade” (manufactured goods to Africa, little green monkey). slaves to the West Indies, sugar to England), and the Cape Verdians shipped salt and live- stock directly to Barbados aboard English ships Grivet, Chlorocebus aethiops (L., 1758) that did not go to Africa as part of the triangle trade (Duncan, 1972). Perhaps the green mon- French: grivet, callitriche keys were introduced to Cape Verde between the Spanish: mono verde 1460s when the Portuguese settlement began, Italian: cercopiteco grigioverde and the 1620s when the British began to settle in German: Grünmeerkatze Barbados (Denham, 1987). In any case, accord- ing to Denham (1987), if the Cape Verde islands Kingdon (1997) and Groves (2001, 2005) served as the source of the monkeys that became recognised the grivet, Chlorocebus aethiops (L., established in the West Indies, it is possible that 1758), as a separate species of the guenon mon- the West Indian populations arose from a much keys, characterised by peculiar morphological pat- more narrowly circumscribed founder popula- terns. Linnaeus (Systema Naturae, 10th ed., 1: 28) tion than has previously been suspected. described this primate on the basis of the exami- ere is also evidence, however, for an earlier nation of specimens from “Ethiopia”. In eff ect, beginning of the exportation of green monkeys the taxon is regarded as an Afrotropical element, beyond their natural distribution, i.e. towards distributed in the savannahs and south Europe. e portrait, for example, of an adult of the Sahara from about 18°-15° N, in Sudan individual of C. sabaeus depicted by the famous east of the White Nile, Eritrea, and Ethiopia Florentine painter Andrea del Sarto in the fres- east of the Rift Valley (Hill, 1966; Haltenorth coes of the Villa Medici of Poggio a Caiano, near & Diller, 1977; Kingdon, 2004; Groves, 2005). Florence, around the end of the second decade In Ethiopia, south of the river Omo, it may cross of the 16th century, testify to an already estab- with C. pygerthrus (Groves, 2001). e record- lished cultural of exporting these living pri- ing of a savannah guenon from the Aïr massif on mates out of Africa, even towards countries very the edge of the Sahara by Bigourdan & Prunier far from the areas of their original distribution (1937), followed by Dekeyser (1950) and Le (Plate 11). e morphological rendering of the Berre (1990), needs confi rmation (Hill, 1966). specimen of green monkey painted by Andrea e latter population was presumably regarded del Sarto is so accurate that the painter was as being related to the form C. aethiops tantalus presumably very familiar with the subject por- Ogilby, 1841. Subsequently, however, Dandelot trayed, and may even have used a live specimen (1959), Kingdon (1997) and Groves (2001) sepa- as a model (cfr. Veracini & Masseti, 2007). is rated C. tantalus from C. aethiops at species level. M. Masseti & E. Bruner 59

Museum collections from the Khartoum Province show that it regularly occurred – even, perhaps, (15°34’N 33°36’E) suggest that grivets survived as a naturalised species – in the Aegean region in Central Sudan until the beginning of the 20th (eastern Mediterranean) in the course of the 2nd century (Peters, 1989). millennium BC. As far as is presently known, According to Hill (1966), in ancient times C. aethiops was imported there from Egypt as a grivets extended their range much further north precious gift, a luxury which the fl ourishing Late than at present. In fact, together with the Barbary Bronze Age palace economy of the Aegean area macaque and the baboon, this was possibly the could aff ord (cf. Masseti, 1997 and 2003b). species of monkey best known in the ancient Mediterranean world. Grivets have been kept Grivets in ancient Egypt as pets since time immemorial and numerous Osborn & Osbornová (1998) are of the examples have been exported out of Africa since opinion that grivets were never actually consid- antiquity. Vague accounts of an East African ered as sacred in ancient Egypt, but were very green monkey have circulated since the days of popular as pets. According to Houlihan (1996) the ancient Greeks, who adopted the term cal- too, they fi gured among the animals that were lithrix (= “beautiful hair”[ cf. Battaglia, 1962; kept as domestic pets. In the artistic produc- Rocci, 1970]) originally introduced by Homer tion, these monkeys were in fact consistently (Hill, 1966). Callitriches were also mentioned depicted as more playful and less serious than by Pliny the Elder who observed that: “Hoc ani- the hamadryas baboons (Arnold, 1995). Very mal negatur vivere in alio quam Aethiopiae quo likely the species was completely absent from gignitur caelo” (= It’s said that this animal can the territory of ancient Egypt, where it was how- survive only in the climate of Ethiopia, where ever frequently imported from far afi eld, from it comes from) (Naturalis historia, VIII, 216). the southern countries such as Nubia and the Although the grivet is not – and perhaps never mythical land of Punt. ese countries roughly has been - a taxon naturally dispersed within correspond respectively to what is now northern the boundaries of the Western Palaearctic, there Sudan and the territories of the so-called Horn exists considerable archaeological evidence to of Africa (Ethiopia and Somalia), where grivets

Fig. 9 - Detail of a painted limestone relief from the mastaba of Ty at Saqqara, Egypt (Fifth Dynasty, 3rd millennium BC).

www.isita-org.com 60 The primates of the western Palaearctic

are still naturally dispersed. Arnold (1995) is, Gabal (Hermopolis) west of the Nile north-west instead, of the opinion that C. aethiops survived of Mallawi, Madinet Gurob and Old Cairo. Two in the wild in ancient Egypt at least up to the grivets were also recorded by Groves (2006) in Middle Kingdom (2134-1785 BC), about the the Ptolemaic-era catacombs al Saqqara, dating same time that the hamadryas baboon also dis- from after 300 B.C. In ancient Egypt, grivets appeared. However, according to Osborn & were not popular only as household pets. In fact, Osbornová (1998), the importation of grivets they were also sent as gifts and traded to vari- from the south, as of other monkeys, doubtless ous parts of the Near East and the Mediterranean began in the earliest dynasties, although they did (McDermott, 1938). not appear in art until much later. e earliest Egyptian portraits of grivets are known from The “blue monkeys” of the Minoan palaces the tombs of the nobles Ty, Ptahotep (Dynasty ere is a considerable amount of evidence to V), and Mereruka (Dynasty VI, 3rd millennium show that grivets were also traded between east- BC), at Saqqara, and from the tomb of Nefermat, ern Africa and the eastern Mediterranean basin in Dynasty IV, at Medum (Osborne & Osbornová, the course of the 2nd millennium BC (Masseti, 1999) (Fig. 9). Images of these monkeys are also 1980, 1997, 2000a, 2003a, 2003b and Masseti, found in the wall-decoration of several tombs of 2006). In the so-called “House of the Frescoes”, the 18th Dynasty (second half of the 2nd mil- west of the palace of Knossos (Crete), for exam- lennium BC), at ebes (Masseti, 1980) (Plate ple, there is a remarkable series of paintings 12 and 13). Grivets are again represented in the showing long-tailed monkeys and medium-sized decoration of the temple of Ramses II, at Beit birds, painted in an intriguing blue colour, set el Wadi south of Aswan (19th Dynasty). As far within a rocky amidst streams, water- as is presently known, mummifi ed grivets were falls and luxuriant vegetation (Late Minoan 1A, listed by Kessler (1989) only from the animal about 1550 BC, Herakleion Museum). Blue mon- cemeteries of the Late Period (end of the 2nd- keys are a recurrent motif in the production of 1st millennium BC) in ebes, Dendera, Tuna el the Aegean Late Bronze Age artists. eir painted

Fig. 10 - Archaeological location of the Minoan sites that provided painted images of “blue mon- keys”. On the upper left, the stone rashly claimed by Poulianos (1972) as the fallacious discovery of a Chlorocebus monkey skull, apparently found on the Aegean island of Santorini (Thera), in the summer of 1966. M. Masseti & E. Bruner 61

images have also been discovered in eran fres- 2004) (Fig. 11). On the other hand, the keeping coes (Late Minoan IA, about 1630 BC Athens, of strange animals by the elite was a widespread National Museum; Plate 14), and in the paintings phenomenon in contemporary Egypt and the from the Pillar Crypt area on the island of Milos Near East (Evely, 1999; Masseti, 2003b). (cf. Morgan, 1990) (Fig. 10). ese painted Morgan (1988) has observed that all the monkeys feature the unmistakable morphologi- scenes in which the “blue monkeys” are depicted cal patterns of the grivet, the exotic primate pre- in Late Bronze Age Aegean art may refl ect con- sumably imported from sub-Saharan Africa into trolled environments, with those apparently the Aegean region through commercial trade set in the wild actually being located in . with Egypt, where it was clearly also regarded as is is suggested essentially by the fact that, very precious (Masseti, 2003b and 2006; see also in view of their value and rarity, these mon- Groves, 2008). e value of the monkeys may keys had to be kept within an area controlled have been related more to their curiosity appeal by humans (Masseti, 1997). Nevertheless, as than to their eff ective economic worth, or pos- already observed by Trantalidou (2000), despite sibly to both (cf. Masseti, 2001). ese monkeys the frequent occurrence of grivets in the artistic were deemed so exotic and important by the production of the Aegean Bronze Age, monkey Aegean Bronze Age that they were not bones are not reported from any sites in the thought of simply as animals, but as creatures Aegean. Poulianos (1972) rashly claimed the invested with the role of intermediary between fallacious discovery of an alleged monkey skull, the human and divine worlds (Marinatos, 1987; apparently found among sea pebbles and rocks Evely, 1999). In the wall-paintings on the upper on the east side of Santorini by the Greek archae- level of the building Xeste 3 of the late Bronze ologist Galanopoulos, in the summer of 1966 Age site of Akrotiri, on the island of Santorini (see Fig. 10). Since it was assumed that this item ( era), a “” has been represented had been covered by the lava that erupted from in a ritual context connected with the off ering the island volcano around the mid 2nd millen- of crocus stigmas, Crocus cartwrightianus, to a nium BC, this was regarded as evidence of the goddess of healing presumably associated with fact “…that the specimen lived during the period saff ron phytotherapy (Ferrence & Bendersky, when Crete was ruled by Minoans and was the

Fig. 11 - In the decoration of the building Xeste 3 of the late Bronze Age site of Akrotiri, on the island of Santorini (Thera), a “blue monkey” was represented in a ritual context connected with the offering of crocus stigmas, Crocus cartwrightianus Herb., to a goddess of healing presumably asso- ciated with saffron phytotherapy.

www.isita-org.com 62 The primates of the western Palaearctic

most powerful state on earth” (Poulianos, 1972). rendering of C. aethiops is so accurate that the Poulianos (1972) even went so far as to indulge in Aegean Bronze Age artists presumably knew the a possible classifi cation of the monkey, referring subject very well, and may even have used live it to the morphology of the representatives of the specimens as models (Masseti, 1980). Beyond Cercopithecidae family, and more specifi cally to a the morphological knowledge displayed, these specimen of “Cercopithecus callitrichus” (cf. Rossi, artists also revealed a specifi c talent in evoking 1968), as grivets were formerly classifi ed (cf. Hill, the behavioural characteristics of the animals 1966). e alleged “skull” was also identifi ed as that they were commissioned to paint (Masseti, belonging to the species of monkey represented 2000, 2003a, 2006). At Knossos, for example, in the murals of Knossos. Subsequently, the aff air grivets are depicted in the course of a raid in a was exposed as a complete fabrication, the article nesting area of the rock doves, Columba livia involved being simply a chunk of lava featuring Gmelin, 1789, which might explain why most a vague resemblance to the exterior of a monkey of the “blue birds” were depicted in fl ight skull (Doumas, 2000). (Cameron, 1968; Masseti, 1997). In fact, one In the Minoan wall-paintings of Knossos, of the primates appears to be eating an egg era and Milos, however, the morphological (Fig. 12). e grivet is in fact often regarded as a species with a marked tendency to raid gar- dens, fi elds and , often becoming a major pest (Haltenorth & Diller, 1977; Despard Estes, 1991). Similar behaviour has been docu- mented in troops of sub-Saharan Africa (Masseti, 2000a, 2003a). Studies carried out on the West- African representative of the guenon monkey taxonomic group – the monkey, C. sabaeus (L., 1766) of northern Senegal – dem- onstrated that these primates can develop the habit of preying upon birds and small mammals, such as columbiformes and murids respectively. Galat & Galat-Luong (1977) observed that this occurred in marginal habitats of the Sahelian zone, particularly in relation to species such as the palm dove, Streptopelia senegalensis (L., 1766), and the Nile rat, Arvichantis niloti- cus (Desmarest, 1822), in periods during which there was an epidemic increase of the latter spe- cies. According to Galat & Galat-Luong (1977), this type of behaviour may arise in the event of competition with rodents, leading C. aethiops to an increased consumption of animal food, including predation upon the Nile rats them- selves. is provides scientifi c verifi cation of the behaviour of the “blue monkeys” in the Knossos wall-painting (Masseti, 2000 and 2006).

Fig. 12 - Detail of the Late Minoan IA wall- painting (c. 1550 BC) from the “House of the Grivets in Europe Frescoes” at Knossos, showing a “blue monkey” Begun in very early times, the exportation of eating an egg from a wild pigeon nest (after grivets beyond the territories of their homeland Cameron, 1968). M. Masseti & E. Bruner 63

has been carried out throughout history. us, European languages, the name given to the grivet for example, pictorial evidence of this monkey is comprises the acknowledgement of an overseas also available among the fi gures in the 12th cen- origin. Eff ectively, the English term guenon and tury fl oor mosaics of the cathedral of Otranto, in the Latin cercopithecus are translated in German Apulia (southern Italy) (Willemsen, 2000). Some by the word Meerkatze, which literally means of the most intriguing images of the species pro- “the cat (Katze) of the sea (Meer)”. duced in Italian art are also found in the works of 14th – 15th century artists such as Pisanello (cf. Cordellier et al., 1996) and/or Giovannino Senegal bushbaby, Galago de’Grassi (cf. Recanati, 2005) (Fig. 13). senegalensis (Geoffroy, 1796) e occurrences of images of C. aethiops in the work of the 16th century Italian naturalist Ulisse French: galago du Sénegal Aldrovandi (V, a., c. 21) - still preserved in the Spanish: gálago del Senegal Biblioteca Universitaria of Bologna (Italy) – pro- Italian: galagone del Senegal, moholi vides evidence of the conspicuous trade contacts German: Steppengalago that still existed in the Middle Ages between Italy, Egypt and eastern North Africa (Capanna It is possible that a relic population of Senegal & Gippoliti, 2007) (Plate 16). Moreover, evi- bushbaby, Galago senegalensis É. Geoff roy, 1796, dence of the popularity of these monkeys in the survived in some remote wadies of the Ennedi western world is underscored by the importance mountainous massif (Chad) up to very recent traditionally attributed to them since Medieval times. is species is a typical Afrotropical ele- times. For example, it is interesting to note how ment whose distribution, according to Hill the grivet was referred to in the pages of the cor- (1953), is limited southwards by the African pus aldrovandianum: “Cercophetecus mas Simia forest belt and to the north by the Sahara. e caudata Gatto Maimone vulgo”. Here it is in Senegal bushbaby has only been reported once fact specifi ed that this long-tailed monkey was from Ennedi by Petit (1937), who observed it in popularly indicated with the name gatto mam- the area of Archei (cf. La Berre, 1990). e region mone. is term was borrowed from the Arabic of Archei (16°20’- 16°50’N 21°50’-21°00’E) is maimūn to indicate the “green monkey”, and located in the western Ennedi, extending over went on to become the Italian mammone, and/ an area of about 18,000 km2. e territory is or gattomammone, a fantastic character in tradi- characterised by the presence of a large “guelta” tional fables, and also the ancient Italian monna - the so-called guelta d’Archei. is is a peculiar and monina (scimmia) (= monkey) (Pellegrini, type of wetland, typical of desert regions, which 1972). Also deriving from this term is the mod- is formed when groundwater in lowland depres- ern Spanish mono (= monkey). However, in other sions rises to the surface, creating permanent

Fig. 13. Some intriguing drawings of grivets produced in Italian art in the 14th-15th centuries.

www.isita-org.com 64 The primates of the western Palaearctic

pools and reservoirs. e peculiar environmental originally distributed in the Holocene bio- conditions of the guelta d’Archei in fact still con- cenoses (M. sylvanus); sent the survival of one of the last populations 2) Borderline species: species of Afrotropical of , Crocodylus niloticus Laurenti, distribution, extending their ranges beyond 1768, known in the Sahara: another biological the boundaries of this biogeographical unit element characteristic of Afrotropical zoogeogra- (P. anubis, P. hamadryas, E. patas, G. senega- phy. Moreover, following Petit (1937), Scortecci lensis); (1940) too mentioned the occurrence of G. sen- 3) Species documented by archaeological evi- egalensis in Ennedi, regarding it as a: “[…] typical dence: taxa documented only by archaeo- element of the Ethiopic zoogeographic region […]”, logical data (C. aethiops); and also positing a hypothetical – albeit never confi rmed - occurrence of the species in Tibesti. 4) Anthropochorous species: species intro- Nevertheless, neither Ellerman & Morrison- duced by humans and subsequently natu- Scott (1951) nor Corbet (1978) include the ralised (C. sabaeus); Senegal bushbaby among the mammalian species In terms of “natural” biogeography, the of Western Palaearctic fauna. Holocenic presence of monkeys in the Western Palaearctic is practically restricted to a single spe- cies, M. sylvanus. Furthermore, while its North Conclusions African range is defi nitely a relic distribution, the European presence in Gibraltar is the result While in many civilisations the role of the of human activity. In fact, we should recall that monkey is traditionally, invested with positive Gibraltar lies beyond the “monkey belts” recog- connotations, in Christian culture (predomi- nised for the zoogeography of primates (Napier nant in the European/Mediterranean countries) & Napier, 1985). the image of this animal has always tended to Most of the other species included in the be negative. Monkeys were frequently taken present study tend to be distributed along the to symbolise the evil or stupid side of human borders of the biogeographical unit in question. beings (e.g., Goves 2008, Herrero Marcos, Olive baboons and patas monkeys are two exam- 2006), which makes all the more intriguing the ples of such distribution, marginal respect to the fact that Darwin’s theory struck the heart of this Western Palaeartic but nonetheless related to its cultural denigration of natural human origins. ecological processes. ese are Afrotropical spe- is cultural bias is very likely related to the lack cies which can be found in the Western Palaearctic of primatological fauna in most of the Western as a result of their desertic or subdesertic habits, Palaearctic territories. Consequently, a study of and the consequent ability to survive in extreme the primates of this biogeographical region is environmental conditions. Some of these groups, rather unusual, since the taxon is hardly repre- such as the Tibesti baboons, appear to have sented. e aim of the present paper is therefore become extinct in very recent times. Others - twofold. Firstly, we have provided an overview of like the baboons in the desert of southern Libya those species originally found in this geographi- - are documented by limited archaeozoologi- cal range. Secondly, we have emphasised the role cal and iconographical evidence, but cannot be of archaeological and ethnozoological records precisely assigned to either a relic or anthropo- in enhancing the biogeographical and historical chorous distribution. A similar comment could knowledge on this group. be made about the current distribution of sacred e species described in this paper can be baboons, in view of their presence in the culture divided into four categories: of ancient Egypt. e Aegean “blue monkeys”, 1) Naturally occurring primates: those taxa instead, are undoubtedly associated with human settlement, and we cannot exclude their possible M. Masseti & E. Bruner 65

naturalisation, albeit for a chronologically lim- Acknowledgements ited period of the protohistoric era. e Cape Verde monkey deserves a special We would like to express our appreciation and mention. Although this species was naturally dis- gratitude to the following friends and colleagues tributed in Western Africa, it was introduced on for their suggestions and assistance in the prepara- Cape Verde possibly in the 17th century, where it tion of this paper: Paolo Agnelli and Saulo Bambi underwent complete naturalisation, to the extent (Natural History Museum of the University of Flor- of being taxonomically described on the basis of ence, Zoological Section), Luca Bachechi, (Istituto specimens from the island of Santiago. Italiano di Preistoria e Protostoria [IIPP], Flor- While the Western Palaearctic has a very ence), Christine Bonnefon (Bibliothèque centrale limited biological record as regards the distri- du Muséum national d’Histoire naturelle, Paris), bution of primates, it has nonetheless played a Stefano Biagetti ( e Italian-Libyan Archaeologi- crucial role in human culture. Over millennia of cal Mission in the Acacus and Messak [Central human transit and commerce, a large amount of Sahara], Sapienza Università di Roma), Andrea zoological information has been encapsulated in Camperio Ciani (Department of General Psychol- “humanistic and artistic media”. e painting, ogy of the University of Padova), Claudio Corridi sculpture and architecture of Europe and the (Florence), Fabrice Cuzin (Marrakesh), Gina Mediterranean off ers a wealth of information on Douglas ( e Linnean Society of London), Elena diff erent aspects of the biodiversity of primates Ferrero (Department of Earth Sciences, University (anthropochorous relocations, historical ranges of Torino), Luca Fiorenza (Senckenberg Museum, and much more besides). One of the major Frankfurt), Caroline Grigson (Institute of Archae- problems in this context is that whenever a pri- ology, University College London), Tarek Jdeidi mate is evoked in literature and/or represented (Zoology Department of the Al-Fateh University, in a work of art or architecture, it tends to be Tripoli), Iannis Machairas (Department of His- generically identifi ed simply “as a monkey”. e tory and Ethnology of the Demokretus Univeristy literary, archaeological and artistic interpreta- of race, Komotini), Iyad Nader (King Khalid tions of the iconography demand a multidisci- Wildlife Research Centre/National Commission plinary approach that exploits both humanistic for Wildlife Conservation and Development, and scientifi c expertise. Here we have attempted Riyadh), Paula Jenkins (Department of Zool- to introduce some of these topics, which clearly ogy of e Natural History Museum, London), call for further study in the appropriate disciplin- Stefania Nosotti (Sezione di Paleontologia of ary context. the Museo Civico di Storia Naturale, Milano), We feel it is important to emphasise the role Claudio Vergano and Paola Lovesio (Associazi- of human populations as “biogeographical vec- one Naturalistica Piemontese), Arturo Morales tors”. Apart from the aforementioned case study (Departamento de Biología, Facultad de Ciencias, of the Cape Verde monkey, and the possible rela- Universidad Autónoma de Madrid) and Cecilia tionships between the North African Veracini (Dipartimento di Biologia Evoluzionis- and baboons (Acacus and ancient Egypt), we tica, Università di Firenze). Spartaco Gippoliti have also referred to the remarkable presence of (Istituto Italiano di Antropologia) and Colin African primates in Neotropical environments. Groves (School of Archaeology and Anthropology. e timing of such processes, and the biologi- Australian national University, Canberra) kindly cal success of certain introductions (i.e., natu- revised a fi rst draft of this manuscript, providing ralisation) raise questions about the proper use of useful comments and suggestions. terms such as “natural” or “artifi cial”, when deal- is research was partially supported by the Istituto ing with expansions of anthropochorous range. Italiano di Antropologia (www.isita-org.com). Eff ectively, this could be regarded as one of the many cases of the joint dispersal of species.

www.isita-org.com 66 The primates of the western Palaearctic

References Ashton E.H. & Zuckerman S. 1950. e infl u- ence of geographic isolation on the skull of the Acidini Luchinat C. 1992. La Grotta degli Ani- green monkey (Cercopithecus aethiops sabaeus). mali. In C. Acidini Luchinat & G. Galletti Part I. Proc. Roy. SocietySer. B, 137: 212-238. (eds): Le ville e i giardini di Castello e Petraia a Ashton E.H. & Zuckerman S. 1951a. e infl u- Firenze, pp. 108-129. Pacini Editore, Ospedal- ence of geographic isolation on the skull of the etto (Pisa). green monkey (Cercopithecus aethiops sabaeus). Afonso-Roque M.M. & Santinho Barata M.C. 2. e cranial dimensions of the St. Kitts and 1992. Filariopsis asper van iel, 1926 (Nem- the African green monkey. Proc. R. Soc.Ser. B, atoda - Metastrongyloidea), parasta dos brôn- 138: 204-213. quios de una primata (Cercopithecus) da ilha Ashton E.H. & Zuckerman S. 1951b. e infl u- de S. Tomé. Garcia da Orta, Sér. Zool., 19: ence of geographic isolation on the skull of the 7-10. green monkey (Cercopithecus aethiops sabaeus). Alhaique F. 2002. Archaeozoology of the funer- 3. e developmental stability of the skulls ary structures. In S. Di Lernia & G. Manzi and teeth of the St. Kitts and the African green (eds): The Archaeology of Libyan Sahara. Vol. monkey. Proc. R. Soc.Ser. B, 138: 213-218. 1. Sand, stones, and bones. The Archaeology of Ashton E.H. & Zuckerman S. 1951c. e infl u- Death in the Wadi Tanezzuft Valley (5000-2000 ence of geographic isolation on the skull of the BP), pp. 181-196. Arid Zone Archaeology green monkey (Cercopithecus aethiops sabaeus). Monograps 3. Università degli Studi di Roma 4. e degree and spread of dental diff erentia- “La Sapienza”/Edizioni All’insegna del Giglio, tion in the St. Kitts green monkey. Proc. R. Soc. Firenze. Ser. B, 138: 354-374. Al-Jumaily M.M. 1998. Review of the mammals Aulagnier A. & evenot M. 1986. Catalogue of the Republic of Yemen. Fauna of Arabia, 17: des mammiferes sauvages du Maroc. Institut 477-502. Scientifi que, Universite Mohammed V/Min- Al-Safadi M.M. 1994. e Hamadryas Babooun, istere de l’Education Nationale, Royame du Papio hamadryas (Linnaeus, 1758) in Yemen Maroc, Rabat. (Mammalia: Primates: Cercopithecidae). Zool- Azzaroli Puccetti M.L. & Zava B. 1988. Nou- ogy in the , 10: 24-36. velles données sur les chiroptères des îles du Anon. 1880. Sketches of natural history in Cap-Vert. Bollettino del Museo Regionale di Sci- Gibraltar. The Field, 10 July 1880. enze Naturali (Torino), 6, 2: 603-615. Aquilué X. 2007. Els animals de companyia Bailey J.F., Henneberg M., Colson I.B., Ciarallo a l’antiguitat. In Aquilué X. & Monturiol J. A., R.E.M. & Sykes B. 1999. Mon- (eds): Animals d’Empúries. La fauna i l’home key business in Pompeii – unique fi nd of a a l’antiguitat. Exposicio, Casa dels Forestals de juvenile Barbary macaque skeleton in Pom- Sant Marti d’Empuries (juliol-setembre 2007). peii identifi ed using osteology and ancient Ajuntament de l’Escala/Museu d’Arqueologia DNA techniques. Mol. Biol. Evol., 16: 1410- de Catalunya-Empuries: 32-35. 1414. Arnold D. 1995. An Egyptian bestiary. The Met- Bailey G., AlSharekh A., Flemming N., Lambeck ropolitan Museum of Art Bulletin, Spring 1995: K., Momber G., Sinclair A. & Vita-Finzi C. 1-64. 2007. Coastal prehistory in the southern Red Ashton E.H. 1959. e infl uence of the geo- Sea Basin, underwater archaeology, and the graphic isolation on the skull of the green Farasan Islands. Proceedings of the Seminar for monkey (Cercopithecus aethiops sabaeus). 5. Arabian Studies, 37: 1-16. e degree and pattern of diff erentiation in Balout L.& Roubet C. 1980. Apports de la the cranial dimensions in the St. Kitts green préhistoire à la stratigraphie et la cronologie au monkey. Proc. R. Soc.Ser. B, 151: 563-583. Sahara et au Maghreb. In M.A. Williams & H. M. Masseti & E. Bruner 67

Faure (eds): The Sahara and the Nile, pp. 163- Historical notes on the early Platyrrhine trade 172. Balkem, Rotterdam. in Italy: the 16th century wall-paintings from Baroni Vannucci A. 1997. Jan Van Der Straet detto Torrechiara castle (Parma). Folia Primatol., 79: Giovanni Stradano fl andrus pictor et inventor. 152. Jandi Sapi Editori, Milano/Roma. Brunner E. 1969. Animals of ancient Egypt. Ani- Battaglia S. (ed). 1962. Grande Dizionario della mals, 11 (12): 559-563. Lingua Italiana. II. Unione Tipografi co-Edi- Buff on G.L. Le Comte de. 1767. Histoire naturelle trice Torinese, Torino. générale et particulière. 14 (Orangs, ). Beck P. & Huard P. 1969. Tibesti carrefour de la L’imprimerie du Roi, Paris. préhistoire saharienne. Arthaud, Paris. Cabrera A. 1914. Fauna Ibérica. Museo Nacional Bellori G. 1931. Le vite dei pittori, scultori et de Ciencias Naturales, Madrid. architetti moderni. Roma. Cabrera A. 1932. Los Mamíferos de Marruecos. Beyhl F.E., Mies B. & Ohm P.1995. Macaronesia Trabajos del Museo Nacional de Ciencias Natu- – A biogeographical puzzle. Bol. Mus. Mun. rales, Serie Zoologica, 57: 1-362. Funchal, Sup. no. 4 : 107-113. Cameron M.A.S. 1968. Unpublished Paintings Bigourdan J. 1950. Présentation des deux pièces from the “House of the Frescoes” at Knossos. zoologiques intéressantes de la collection de The Annual of the British School of Archaeology IFAN. C.R. Première Conf. Internat. African. at Athens, 63: 1-31. De l’Ouest, I, Dakar. Camperio Ciani A. 1986. Origine, evoluzione, Bigourdan J. & Prunier R. 1937. Les mammifères speciazione ed ecologia del genere Macaca in sauvages de ‘Ouest Africain et leur mileu. Rud- relazione alle vicende geoclimatiche del Qua- der, Montrouge. ternario. Studi per l’Ecologia del Quaternario, Boessneck J. 198. Die Tierwelt des Alten Ägypten. 8: 9-32. München. Camperio Ciani A., Palentini L., Arahou M., Borzatti von Löwenstern E. 1982. Quando il Martinoli L., Capiluppi C. & Mouna M. Sahara era verde. Paravia, Torino. 2005. Population decline of Macaca sylvanus Borzatti von Löwestern E. & Masseti M. 1991. in the Middle Atlas of Morocco. Biol. Conserv., On the twisted –horn antelopes carved in the 121: 635-641. rocks of some wadies of the Jordan southern Capanna E. & Gippoliti S. 2007. I “Quadru- desert. Studi per l’Ecologia del Quaternario, 13: pedi Vivipari”. In A. Alessandrini & A. Cere- 143-148. gato (eds): Natura picta. Ulisse Aldrovandi, pp. Bourbon S. de. 1932. De l’Algérie au Tchad 83-87. Editrice Compositori, Bologna. par le Hoggar et l’Aïr. Bull. Com. Et. Hist. Sc. Carl L. & Petit J. 1954. La ville de sel (du Hoggar A.O.F., 13: 1-183. au Tibesti). René Julliard, Paris. Bourbon H.R.H. Prince Sixte of. 1933. Great Cassoli P.F. & Durante S. 1974. La fauna del Ti- routes of the Sahara, past and present. Royal n-Torha (Acacus, Libia). Origini, 8: 159-161. Geographical Society, London. Castells Á. & Mayo M. 1993. Guía de los Bourguignat J.R.. 1870. Histoire du Djebel-Thaya mamíferos en libertad de España y Portugal. et des ossements fossiles recueillés dans la grande Ediciones Pirámide, Madrid. caverne de la Mosqueé. Paris. Cavicchio P. & Friedrich K.G. 2006. Igiene e Bousquet B. 1992. Guide des parcs nationaux sicurezza nella gestione delle collezioni prima- d’Afrique. Delachaux et Niestlé, Paris. tologiche. In E. Bruner & S. Gippoliti (eds): Le Brown L. 1965. Africa, a natural history. Chanti- Collezioni Primatologiche Italiane, pp. 53-64. cleer Press Edition, New York. Istituto Italiano di Antropologia, Roma. Brack M. 1987. Agents transmissible from Chapman N.G. & Chapman D.I. 1980. e dis- to man. Springer-Verlag, Berlin. tribution of fallow deer: a worldwide review. Bruner E., Masseti M. & Veracini C. 2008. Mammal Review, 10: 61-138.

www.isita-org.com 68 The primates of the western Palaearctic

Charco J. 1999. El bosque mediterráneo en le norte Palaeoenvironment and prehistory in south- de África. Agencia Española de Cooperación western Fezzan (Libyan Sahara), pp. 89-94. Internacional. Madrid. C.I.R.S.A. Centro Interuniversitario di Ricerca Chiarini M. 1977. e Decoration of Palazzo per le Civiltà e l’Ambiente del Sahara Antico. Pitti in the Seventeenth and Eighteenth Cen- Edizioni All’Insegna del Giglio, Firenze. turies. Apollo: 178-202. C.N.R. Quaderni di Geodinamica Alpina e Churcher C.S., Kleindienst M.R., Wiseman M.F. Quaternaria, 7. & McDonald M.M.A. 1997. e Quaternary Cortés J.E., Finlayson J.C., Mosquera M.A. & faunas of Dakhla Oasis, Western Desert of Garcia E.F.J. 1980. The birds of Gibraltar. Egypt. Abstracts of the Second Dakhleh Oasis Gibraltar Booshop, Gibraltar. Project Research Seminar.June, 16-20, 1997, Coutinho Saraviva A. 1961. I. Biogeografi a. In A. 1-3. Royal Ontario Museum and University of Coutinho Saraviva (ed): “Conspectus” de ento- Toronto, Toronto. mologia cabo-verdiana: 19-34. Junta de Investi- Ciarallo A. & De Carolis E. 1999. Pompeii. Life gações do Ultramar, Lisbon. Estudos, Ensaios e in a Roman Town. Homo faber. Natura, Scienza Documentos, 83. e Tecnica nell’antica Pompei. Electa, Milano. Cramp S. (ed). 1977. Handbook of the Birds of Clarke T.H. 1986. The Rhinoceros from Dürer Europe, the Middle East and North Africa. to Stubbs 1515-1799. Sotheby’s Publication, e Birds of the Western Palearctic. Volume London. I. Ostrich to Ducks. Oxford University Press, Cloudsley- ompson J.L. 1984. Key Environ- Oxford/London/New York. ments: Sahara Desert. Pergamon Press, Elms- Crooks K.R. & Rasmussen D.R. 1991. e rela- ford, New York. tionships between and grooming in Colyer J.F. 1948a. e green monkey of St. Kitts. the Tanaxpillo colony of stumptail macaques Dental instability. Br. Med. J., 1: 1202. (Macaca arctoides). Laboratory Primate News- Colyer, J.F. 1948b. Variations of the teeth of the letter, 30: 2-5. green monkey of St. Kitts. Proc. R. Soc. Med., Cuzin F. 1996. Répartition actuelle et statut des 61: 845-848. grands mammiferes sauvages du Maroc (Pri- Corbet G.B. 1978. The mammals of the Palaearc- mates, Carnivores, Artiodactyles). Mammalia, tic Region: a taxonomic review. British Museum 60: 101-124. (Natural History)/Cornell University Press, Cuzin F. 2003. La régression des grands mam- London and Ithaca. mifères terrestres du Maroc méridional au Corbet G.B. & Hill J.E. 1992. The mammals of course du XXe siècle. Naturalia maroccana, 1: the Indomalayan Region: a systematic review. 93-99. Natural History Museum Publications/Oxford Dacos N. 1969. Présents américains a la Renais- University Press, Oxford. sance. L’assimiliation de l’exotisme. Gazette des Cordellier D., Lanfranc de Pantou C., Moench Beaux-Arts, 105 Sup.: 58-64. E., Rossi F., Toscano G. & de Turckheim-Pey S. Dacos N. 1977. Le logge di Raffaello. Maestro e 1996. Pisanello. Le peintre aux sept vertus. Editions bottega di fronte all’antico. Istituto Poligrafi co de la Réunion des musées nationaux, Paris. dello Stato, Roma. Corridi C. 1992. L’archeozoologia e le faune olo- Dalloni M. 1935. Mission du Tibesti. Mam- ceniche del Sahara. In M. Lupacciolu (ed.): mifères. Mém. Acad. Sci. Paris, 62: 41-43. Arte e culture del Sahara preistorico, pp. 73-76. Dandelot P. 1959. Note sur la classifi cation des Casa Editrice Quasar, Roma. Cercopithèques du groupe aethiops. Mamma- Corridi C. 1998. Faunal remains from Holocene lia, 23: 357-368. archaeological sites of the Tadrart Acacus and Deag J.M. 1977. e status of the Barbary surroundings (Libyan Sahara). In M. Cre- macaque Macaca sylvanus in captivity and fac- maschi & S. Di Lernia (eds): Wadi Teshuinat. tors infl uencing its distribution in the wild. M. Masseti & E. Bruner 69

In S.H. Prince Rainer III of Monaco & G.H. Chicago Press, Chicago. Bourne (eds): Primate Conservation, pp. 267- Duveyrier H. 1864. L’exploration du Sahara. Les 287. Academic Press, New York. Touaregs du Nord. Challamel, Paris. Deag J.M. & Crook J.H. 1971. Social behaviour Edwards G. 1758-1764. Gleanings of natural his- and “agonistic buff ering” in the wild Barbary tory. Royal College of Physicians, London. macaque Macaca sylvana L. Folia Primatol., Ellermann J.R. & Morrison-Scott T.C.S. 1951. 15: 183-200. Checklist of Palaearctic and Indian mammals Dekeyser P.L. 1950. Mammifères. Contribution 1758 to 1946. British Museum (Natural His- al’Étude de l’Aïr. Mémoires de l’Institut Français tory), London. d’Afrique Noir, pp. 388-425. Librairie Larose, Elliot D.G. 1913. A review of the Primates. Paris. Amer. Mus. Nat. Hist., 11: 147. Dekeyser P.L. 1952. A propos de la tête osseuse Erman A. 1894. Life in ancient Egypt. 1971 edi- d’un cinocéphale du Tibesti. Bull. Institut tion. New York. Française Afrique Noire,Dakar, 14: 537-544. Evslin B. 1988. Gods, demigods and demons. I.B. Dekeyser P.L. 1955. Les mammifères de l’Afrique Tauris & Co. Ltd, London. noire française (2e edition). Institut français Fa J.E. 1981. Apes on the rock. Oryx, 16: d’Afrique noire, Dakar. 73-76. Dekeyser P.-L. & Derivot J. 1959. La vie animale Fa J.E. 1984. Habitat distribution and prefer- au Sahara. Librairie Armand Colin, Paris. ences in the Barbary macaque. Int. J. Prima- Delson E. 1980. Fossil macaques, phyletic rela- tol., 5: 273-286. tionships and a scenario of deployment. In Fa J.E. 1999. Macaca sylvanus (L., 1758). In A.J. D.G. Lindburg (ed.): The Macaques: Studies in Mitchell-Jones, G. Amori, W. Bogdanowicz, B. Ecology, Behaviour and Evolution: 10-30. Van Kryštufek, P.J.H Reijnders, F. Spitzenberger, M. Nostrand Reinhold, New York. Stubbe, J. B. M. issen, V. Vohralik & J. Zima, Denham W.W. 1987. Green monkey migration 1999 (eds): The Atlas of the European mammals, revisited. Review of West Indian Green Monkeys: pp. 158-159. Academic Press, London. Problems in Historical Biogeography. Karger, Fa J.E., Taub D.M., Menard N. & Stewart P. Basel. 1984. e distribution and current status of De Oliveira Boléo J. 1939. O descobrimento e os the Barbary macaque in North Africa. In J.E. descobridores das ilhas do Cabo Verde. Grupo Fa (ed): The Barbary macaque: a case study in Portugês da História das Ciências, Lisbon. conservation, pp. 79-111. Plenum Press, New De Vasconcelos E. 1920. Colónias Portuguesas. York. Arquipelago de Cabo Verde. Lisbon. Fenoyl R. de (ed) 1980. Histoire Naturelle de Di Lernia S. & Manzi G. 2002. Sand, Stones, l’Egypte par Prosper Alpin 1581-1584. Institut and Bones. The archaeology of death in the Wadi Français d’Archéologie Orientale, Paris. Tanezzuft Valley (5000-2000 bp). e Archaeol- Ferrence S.C. & Bendersky G. 2004. erapy ogy of Libyan Sahara, Volume I. AZA Mono- with saff ron and the goddess at era. Persp. graphs 3. All’Insegna del Giglio, Florence. Biol. Med., 47: 199-226. Doumas C. 2000. Discussion. In K. Trantalidou: Flamand G.B.M. 1902. Sur l’utilisation comme Animal bones and animal representations at Late instruments néolithiques de coquilles fossiles Bronze Age Akrotiri. In S. Sherratt (ed): The à taille intentionnelle (littoral du Nord Afri- wall paintings of Thera. Vol. III, pp. 534-735. cain). Assoc. Fr. Avanc. Sci., Ajaccio, 2, 1901 era Foundation-Petros M. Nomikos and the (1902): 729-734. era Foundation, Athens. Fontoura da Costa A. 1937. Deambulações da Duncan T.B. 1972. Atlantic islands: Madeira, the ganda de Modafar, rei de Cambia, da 1514 a Azores, and the Cape Verdes in the seventeenth 1516. República Portuguesa, Ministério das century commerce and navigation. University of Colónias, Lisbon.

www.isita-org.com 70 The primates of the western Palaearctic

Fooden J. 1964. Rhesus and crab-eating Gautier A. 1987b. e archaeozoological macaques: intergradation in ailand. Science, sequence of the Acacus. In B.E. Barich (ed): 143: 363-365. Archaeology and environment in the Libyan Fooden J. 1976. Provisional classifi cation and Sahara. The excavations in the Tadrart Acacus, key to living species of macaques. Folia Prima- 1978-1983, pp. 283-312. BAR International tol., 25: 225-236. Series, 368. Fooden J. 1980. Classifi cation and distribution Gautier A. 1993. Mammifères holocènes du Sahara of living macaques. In D.G. Lindburg (ed): d’après l’art rupestre et l’archaèozoologie. In The macaques. Studies in ecology, behaviour and G. Callegari (ed): L’arte e l’ambiente del Sahara evolution, pp. 1-8. Van Nostrand Reinhold, preistorico : dati e interpretazioni, pp. 261-267. New York. Memorie della Società Italiana di Scienze Nat- Fooden J., Quan Guo-qiang, Wang Zong- urali e del Museo Civico di Storia Naturale di ren & Wang Ying-xiang. 1985. e stump- Milano, 26, 2. tail macaques of China. Am. J. Primatol., 8: Gautier A. & Van Neer W. 1977. Prehistoric 11-30. fauna from Ti-n-Torha (Tadrart Acacus, Fooden J. 2000. Systematic review of rhesus Libya). Origini, 9: 87-127. macaque, Macaca mulatta (Zimmermann, Goldsmith O. 1840. A history of th earth and ani- 1780). Fieldiana Zoology, 96: 1-180. mated nature. Blackie and Son, Glasgow. Fooden J. 2007. Systematic Review of the Bar- Goudsmith J. & Brandon-Jones D. 1999. Mum- bary Macaque, Macaca sylvanus (Linnaeus, mies of olive baboons and Barbary macaques 1758). Fieldiana Zoology, 113: 1-60. in the Baboon Catacomb of the Sacred Ani- Frade F. 1956. Aves e mamiferos da ilhas de São mal Necropolis at North Saqqara. J. Egypt. Tomé e do Principe – Notas de sistematica e Archaeol., 85: 45-53. de protecção à fauna. CCTA/CSA Conferencia Gonzales Henriquez M.N., Rodrigo Perez J.D. & Internacional dos Africanistes Ocidentais, 6a Suarez C. 1986. Flora y vegetación del Archip- Sess°o. S. Tomè: 137-149. iélago Canario. Cedirca s.L. edición regional Fryer J. 1909. A new account of East and canaria, Las Palmas de Gran Canaria (Canary Persia (edited by W. Crooke). e Hakluyt islands). Society, London. Groves C.P. 2001. Primate taxonomy. Smithso- Funaioli U. 1971. Guida breve dei mammif- nian Institution Press, Washington DC. eri della Somalia. Istituto Agronomico per Groves C.P. 2005. Order Primates.In D.E. Wil- l’Oltremare, Florence. son & D.-A.M Reeder. (eds): Mammal species of Galat G. & Galat-Luong A. 1977. Demographie the world. A taxonomic and geographic reference. et regime alimentaire d’une troupe deCercop- ird edition, Volume 1, pp. 111-184. e ithecus aethiops sabaeus en habitat marginal au John Hopkins University Press, Baltimore. nord Senegal. La Terre et la Vie, 31: 557-577. Groves C., 2006. From Calpe to Catanduanes: Garcia J. 1979. The famous rock apes of Gibral- Babewynes, apes, marmesettes and othere tar. Mediterranean Sun Publishinf Co Ltd, diverse bestes. In Hodges J.K. & Cortes J. Gibraltar. (eds): The Barbary Macaque: Biology, Manage- Gasperetti J. 1981. Birds of Saudi Arabia. A note ment and Conservation, pp. 1-15. Nottingham on Arabian ornithology – Two endangered spe- University Press. cies. Fauna of Saudi Arabia, 3: 435-440. Groves C. 2008. Extended family: long lost Gautier A. 1987a. Archaeozoology of the Bir cousins. A personal look at the history of pri- Kiseiba region, Eastern Sahara. In F. Wendorf, matology. Conservation International, Arling- R. Schild & Close A.E. (eds): Cattle-Keepers of ton (USA). the Eastern Sahara, pp. 49-76. S.M.U. Press, Guardia J. & Maragall M., 2004. Periodització Dallas. del jaciment de les Colomines, Zona A (Llívia, M. Masseti & E. Bruner 71

Cerdanya). Fase constructives i seqüencia Hill W.C. O. 1970. Primates. Comparative anat- cronologica. Setenes Jornades d’Arqueologia de omy and taxonomy. Volume 8. Cynopithecinae. les comarques de Girona (La Bisbal d’Empordà, Edinburgh University Press, Edinburgh. 4 i 5 de juny de 2004), Girona, 2004: 247- Ho C.H. & Chiu S.T. (eds). 1983. Nature Con- 252. servation in China. Hong Kong Museum. Guardia J., Maragall M., Mercadal O., Olesti Honh Kong. O., Galbany J. & Nadal J. 2005. Enterrament Hodges J.K. & Cortes J. 2006- e Barbary d’època tardoromana corresponnent a un Macaque: Biology, Management and Conser- macaco amb aixovar al jaciment de les Colo- vation. Nottingham University Press, UK. mines (Llívia). Ceretania, 4: 66-106. Houlihan P.F. 1996. The animals world of the Guest E. & Al-Rawi A. 1966. Introduction to pharaohs. ames and Hudson, New York. the fl ora. Flora of Iraq, 1. Ministry of Agricul- Jardine W. 1833. The naturalist’s library. Vol. 1 ture of the Republic of Iraq/University Press (mammalia) monkeys. Lizars and Stirling and Glasgow, Baghdad: pp.1-214. Kenny, Edinburgh. Hacid M. 1992. Les pierres écrites de l’Atlas saha- Jobst W., Erdal B. & Gurtner C. 1997. Istanbul. rien (El-Hadira el-Mektouba). ENAG, Alger. The great palace mosaic. Arkeoloji ve Sanat Hall D.N., Williams M.A.J., Desmond Clark Yayinlari, Istanbul. J., Warren A., Bradley P. & Beighton P. 1971. Jolly D.W. & Rasmussen D.R. 1991. Use of the e British expedition to the Aïr mountains. islands for propagation of Geographical Journal, 137, 4: 30-36. and ecotourism. American Journal of Primatol- Haltenorh T. & Diller H. 1977. Säugetiere Afri- ogy, 24: 110. kas und Madagaskars. BLV Verlagsgesellschaft Johnsgard P. 1988. The quails, partridges, and mbH, München. francolins of the world. Oxford University Press, Harrison D.L. 1964. The mammals of Arabia. Oxford/New York/Tokyo. Vol. 1. Ernest Benn Ltd, London. Jones T., Erhardt C.L., Butynski T.M., Daven- Harrison D.L. 1972. The Mammals of Arabia. port T.R.B., Mpunga N.E., Machaga S.J. & Vol. III. Lagomorpha, Rodentia. Ernest Benn De Luca D.W. 2005. e Highland Manga- Ltd, London. bey, Lophocebus : a new species of Afri- Harrison D.L. & Bates P.J.J. 1991. The Mam- can monkey. Science, 5752: 1161-1164. mals of Arabia. Harrison Zoological Museum, Keimer L. 1939. Pavian and Dum Palme. Mit- Sevenoaks (Kent). teilungen des Deutschen archäologischen Insti- Hausfater G. 1974. History of three little-known tutes, Abtailung Kairo, 8: 42-45. populations of macaques. Labora- Kessler D. 1989. Die heiligen Tiere und der König. tory Primate Newsletter, 13, 1: 16-18. Vil. I. Beiträge zu Organization, Kult und Theolo- Hazevoet C.J. 1995. The birds of the Cape Verde gie der spätzeintlichen Tierfriedhöfe. . islands. British Ornithologist’s Union, Tring. King A. 2002. Mammals. Evidence from wall Hemprich F.G. & Ehrenberg C.G. 1829. Beo- paintings, sculpture, mosaics, faunal remains, bachtungen über die Aff en-Rten in Sennaar, and ancient literary sources. In W. Feemster Cordofan und Arabien. Verh. Ges. naturf. Fre- Jashemski & F.G. Meyer (eds): The natural unde Berlin, 6: 406-408. history of Pompeii, pp. 401-450. Cambridge Hill W.C. O. 1953.Primates.Comparative anat- University Press, Cambridge. omy and taxonomy. Volume 1. Strepsirini. Edin- Kingdon J. 1997. The Kingdon field guide to Afri- burgh University Press, Edinburgh. can mammals. Nature World Academic Press. Hill W.C. O. 1966. Primates. Comparative Harcourt Brace & Company, San Diego. and taxonomy. Volume 6. Catarrhini Kowalski K. & Rzebik-Kowalska B. 1991. Cercopithecoidea . Edinburgh Mammals of Algeria. Ossolineum, Wroclaw/ University Press, Edinburgh. Varsavia/Cracovia.

www.isita-org.com 72 The primates of the western Palaearctic

Kummer H., Banaja A.A., Abo-Khatwa A.N. MacDonald J. 1965. Almost human. Phyla, New & Ghandour A.M. 1981. Mammals of Suadi York. Arabia. Primates. A Survey of Hamadryas Macdonald A.A. & Yang L.N. 1997. Chi- Baboons in Saudi Arabia. Fauna of Saudi Ara- nese sources suggest early knowledge of the bia, 3: 441-471. ‘unknown’ ungulate (Pseudoryx spiralis) from Kunkel G. 1980. Die Kanarischen Inseln und ihre and Cambodia. J. Zool. London, 241: Pflanzenwelt. Gustav Fischer Verlag, Stuttgard/ 523-526. New York. McDermott W. 1938. e ape in antiquity. Johns Kunz J. 1988. Une site à peinture pariétales de Hopkins Studies in Archaeology, 27: xii + 338. l’Oued Tirenhaut-Tassili n’Ajjer. Sahara, 1 : Machairas I., Camperio-Ciani A. & Sgardelis 95-96. S. 1999. Habitat use by Macaca sylvanus in Kurtén B. 1968. Pleistocene mammals of Europe. two localities of the Moroccan Middle Atlas. Weidenfeld and Nicolson, London. Abstracts of the 8th International Congress on the Lakhdar A., Fartouat J.P., evenot M. & Zoogeography and Ecology of Greece and Adja- Richert N. 1975. Faune du Maroc. Les mam- cent Countries: 88.. Kavala, 17-21 May 1999. mifères. Institut d’études et de recherches pour Martin R.D. & Segesser F. von. 1996. Fragmen- l’arabisation, Rabat. tation of natural populations, genetics and Lavauden L. 1926. Les Vertébrés du Sahara. Élé- conservation biology. Almoraima, 15: 311- ments de zoologie saharienne. Imprimerie Albert 326. Guénard, Tunis. Martin R.P. & Hirschfeld E. 1998. Comments Lavauden L. 1927. Les Forêts du Sahara. Revue on the limits of the Western Palearctic in Iran des Eaux et Forêts, juin-juillet 1927: 24-36. and the Arabian Peninsula. Sandgrouse, 20: Lavauden L. 1930. Une nouvelle perdrix des 108-134. montagnes du Sahara central. Alauda, Etutes et Mason M. 1936. Présence du singe rouge (Eryth- Notes ornithologiques, 3-4: 1-5. rocens patas) au nord du Wadi Hawa- 300 Lawson Handley L.J., Hammond R.L., Emaresi miles from the nearest open water. Geographi- G., Reber A. & Perrin N. 2006. Low Y chro- cal Journal, 216-217. mosome variation in Saudi-Arabian hamad- Masseti M. 1980. Le scimmie azzurre. La fauna ryas baboons (Papio hamadryas hamadryas). etiopica degli aff reschi minoici di Santorino Heredity, 96: 298-303. ( era). Mondo archeologico, 51: 32-37. Le Berre M. 1990. Faune du Sahara. 2. Mam- Masseti M. 1991. Dalla “turata della Gran’Bestie” mifères. Terres africaines. Lechevalier – R. allo “stanzone” degli agrumi: splendore e deca- Chabaud, Paris. denza dei serragli faunistici del Giardino di Lever C. 1985. Naturalized mammals of the world. Boboli. In C. Acidini Luchinat & E.Garbero Longman, London and New York. Zorzi (eds): Boboli 90. Atti del Convegno Liverani M. 1994. Guerra e diplomazia nell’Antico Internazionale, pp. 323-337. Edifi r-Edizioni Oriente 1600-1100 A.C. Editori Laterza, Firenze, Florence. Roma-Bari. Masseti M. 1996. e postglacial diff usion of the Lord Lilford. 1866. Notes on the ornithology of genus Dama Frisch, 1775, in the Mediterra- Spain. Ibis, 2: 173-187, 384. nean region. Suppl. Ric. Biol. Selvaggina, 25: Loy J.D. 1974. Changes in facial color associ- 7-29. ated with pregnancy in patas monkeys. Folia Masseti M. 1997. Representation of birds in Primatol., 22: 251.257. Minoan art. Int. J. Osteoarchaeol., 7: 354- Lynn C.J. 1997. Excavations at Navan Fort 363:. 1961-71, County Armagh. In D.M. Water- Masseti M. 2000. Did the study of ethology man (ed): Northern Ireland Archaeological begin in Crete 4000 years ago?. Ethology Ecol- Monographs, 3. Stationery Offi ce, Belfast. ogy & Evolution, 1: 89-96. M. Masseti & E. Bruner 73

Masseti M. 2002. Uomini e (non solo) topi. Mohammed-Ali A.S. 1981. Archaeological sur- Gli animali domestici e la fauna antropocora. vey in the Wadi Hawar basin. Curr. Anthropol., Firenze University Press/Università degli Studi 22, 2: 176-178. di Firenze, Florence. Morales A. 1994. Earliest genets in Europe. Masseti M. 2003a. Taxonomic and behavioural Nature, 370: 512-513. aspects of the representation of mammals in Morales A. & Rofes J. 2008. Early evidence for Aegean Bronze Age art.) In E. Kotjabopoulou, the Algerian hedgehog in Europe. J. Zool. 274: Y. Hamilakis, P. Halstead, C. Gamble & P. 9-12. Elefanti (eds): Zooarchaeology in Greece. Recent Morales J.C. & Melnick D.J. 1998. Phylogenetic advances, pp. 267-275. British School at Ath- relationships of the macaques (Cercopitheci- ens Studies, 9. dae: Macaca), as revealed by high-resolution Masseti M. 2003b. Exploitation of fauna in the restriction site mapping of mithocondrial Near East during ancient historical times: ribosomal genes. J. Hum. Evol., 34: 1-23. observations on the mammalian species repre- Morgan L. 1988. The miniature Wall-Paintings sented in Assyrian art. In C. Albore Livadie & from Thera: a Study in Aegean Culture and F. Ortolani (eds): Variazioni climatico-ambien- Iconography. Cambridge University Press, tali e impatto sull’uomo nell’area circum-medi- Cambridge. terranea durante l’Olocene, pp. 375-390. Cen- Mori F. 1960. Arte Preistorica del Sahara Libico. tro Universitario Europeo per i Beni Culturali, De Luca, Roma. Ravello. Edipuglia, Bari. Mori F. 1965. Tadrart Acacus. Arte rupestre e cul- Masseti M. 2006. Did the study of primatology ture del Sahara preistorico. Einaudi, Torino. begin on the island of Crete (Greece) 4000 Mori F. 2000. Le grandi civiltà del Sahara antico. years ago? Folia Primatol., 77: 267-301. Bollati Boringhieri, Torino. Masseti M. 2008. Sculptures of mammals in Muzio C. (ed.). 1925. Isole africane. Mundus, the Grotta degli Animali of the Villa Medici Monografi a 49a. Casa Editrice Sonzogno, di Castello, Florence, Italy: a stone menagerie. Milano. Archives of Natural History, 35: 100-104. Muzzolini A. 1989. New fi nds of late Round Meankin B. 1901. The land of the Moors. Darf Head paintings in northern Tassili and the Publishers Ltd, London (1986). break of the “Postneolithic Arid Phase”. Mehlman P. 1989. Comparative density, demog- Nyame Akuma. Bulletin of the Society of Afri- raphy, and ranging behavior of Barbary canist Archaeologists, 31: 2-8. macaques (Macaca sylvanus) in marginal and Nader I.A. 1990. Checklist of the mammals of prime conifer habitats. Int. J. Primatol., 10: Arabia. Fauna of Saudi Arabia, 11: 329-381. 269-292. Napier J.R. & Napier P.H. 1967. A handbook of Ménard N., Vallet D. & Gautier-Hion A. 1985. living primates. Academic Press, London. Démographie et reproduction de Macaca syl- Napier J.R. & Napier P.H. 1985. The natural vanus dans diff érents habitats en Algérie. Int. history of the primates. MIT Press, Cambridge J. Primatol., 4 : 65-81. (Massachusetts). Miller G.S. 1942. Zoological results of the Napier P.H. 1981. Catalogue of Primates in the George Vanderbilt Sumatran Expedition, British Museum (Natural History) and elsewhere 1936/39. Part V. Mammals collected by Fred- in the British isles. Part 2. Family Cercopitheci- erick A. Ulmer, Jr. on Sumatra and Nias. Proc. dae, subfamily Cercopithecinae. British Museum Acad. Nat. Sci. Philad., 94: 107-165. (Natural History), London. Modolo L., Salzburger W. & Martin R.D. 2005. Naurois R. de. 1969. Notes brèves sl l’Avifaune de Phylogeography of Barbary macaques (Macaca l’Archipel du Cap Vert. Faunistique, endèmisme, sylvanus) and the origin of the Gibraltar colony. écologie. Bull. Inst. Fondam. D’Afrique Noire Proc. Natl. Acad. Sci USA, 102: 7392-7397. (IFAN), 31, serie A, 1: 143-218.

www.isita-org.com 74 The primates of the western Palaearctic

Naurois R. de. 1994. Les oiseaux de l’archipel du Bulletin de la Société d’Histoire Naturelle de Cap Vert. Instituto de Investigação Cientí- l’Afrique du Nord, 28 : 392-405. fi ca Tropical, Ministério do Planeamento e Petragnani E. 1928. Il Sahara tripolitano. Collezi- da Administração do Territorio, Secreteria de one di opere e di monografi e a cura del Min- Estrado da Ciência e Tecnologia, Lisbon. istero delle Colonie, 3. Sindacato Italiano Arti Oates J.F. 1996. African primates: status, survey Grafi che, Roma. and conservation action plan. Revised edition. Pocock R.I. 1907. A monographic revision of IUCN, Gland (Switzerland). the monkeys of the genus Cercopithecus. Proc. Ogilby W. 1838. The Menageries; the Natural Zool. Soc. London: 677-746. History of Monkeys, Opossum and Lemurs. C. Pocock R.I. 1932. e Rhesus macaques (Macaca Knight, London. mulatta). J. Bomb. Nat. Hist. Soc., 35: 530- O’Keefe R.T. & Lifshitz K. 2006. A behavioural 551. profi le for stumptail macaques (Macaca arc- Poulianos A.N. 1972. e discovery of the fi rst toides). Primates, 26, 2: 143-160. victim of era’s Bronze Age eruption. Archae- Osborn D.J. & Osbornová J. 1998. The mam- ology, 25, 3: 229-230. mals of ancient Egypt. Aris & Phillips Ltd, Pritchard J.M. 1979. Landform and landscape in Warminster (England). Africa. Edward Arnold, London. Palagi E. 1998. La Galleria dei Primati. Museo di Rabb M.S. 1968. e sacred baboon. Chicago Storia Naturale e del Territorio dell’Università Zoological Society, 34: 2-4. di Pisa/Edizioni ETS, Pisa. Recanati M.G. 2005. Il Taccuino di Giovannino Pandrea I., Apetrei C., Dufour J., Dillon N., de’Grassi. Franco Maria Ricci, 8: 108-132. Barbercheck J., Metzger M., Jacquelin B., Renaud P. 1976. Ile Maurice. Les Editions du Bohm R., Marx P.A., Barre-Sinoussi F., Hirsch Pacifi que, Papeete (Tahiti). V.M., Müller-Trutwin M.C., Lackner A.A. & Ribeiro H., da Cunha Ramos H., Antunes Capela Veazey R.S. 2006. immunodefi ciency R. & Alves Pires C. 1980. Os mosquitos de virus SIVagm.sab infection of Caribbean Afri- Cabo Verde. Junta de Investigações Cientifi cas can green monkeys: a new model for the study do Ultramar, Lisbon. Estudios, Ensaios e Docu- of SIV pathogenesisin natural hosts. J. Virol., mentos, 135: 1-141. 80: 4858-4867. Richard A.F., Goldstein S.J. & Dewar R.E. 1989. Paolucci A. 2000. The animals of Giambologna. Weed Macaques: e Evolutionary Implica- Italian Ministry for Cultural Assets and Activi- tions of Macaque Feeding Ecology. Int. J. Pri- ties, Fine Arts and Historical Assets Service of matol., 10: 569-591. Florence, Pistoia, and Prato/Giunti Gruppo Righetti R. 2004. Capo Verde. Ulysse Network Editoriale, Florence. srl, Milano. Payne R. 2003. A Report from birdtours.co.uk. Rocci L. 1970. Vocabolario Greco-Italiano. Soci- Cape Verde Islands, 3rd to 16th March 2003. età Editrice Dante Alighieri, Milano, Rome, Birdwatching Trip Reports, Birdtours.co.uk: 4. Naples and Città di Castello/Società Editrice Pellegrini G.B. 1972. Gli arabismi nelle lingue S. Lapi, Città di Castello. neolatine con speciale riguardo all’Italia. I. Paid- Rode P. 1938. Catalogue des types des mammif- eia Editrice, Brescia. eres au Museum nationale d’histoire naturelle. Peters J. 1989. Faunal remains and environmen- Bull. Mus. Paris, 2, ser. 10: 202-251. tal change in Central and Eastern Sudan from Rolfe W.D.I. & Grigson C. 2006. Stubbs’s “ Terminal Pleistocene to Middle Holocene and albino hamadryas baboon” in conjectural times. Acad. Wetenschap. Lett. Schon. Kunst. historical context. Arch. Nat. Hist., 33: 18-41. België, 51, 4: 123-148. Rook L., Martínez-Navarro B. & Clark Howell Petit G. 1937. Matériaux de la mission d’étude de F. 2004. Occurrence of sp. in la biologie des acridiens. Vertébrés de l’Ennedi. the Late Villafranchian of Southern Italy and M. Masseti & E. Bruner 75

implication for Early Pleistocene “out of Africa” Soleilhavoup F. 2003. Art préhistorique de l’Atlas dispersals. J. Hum. Evolut., 47: 267-277. saharien. Pilote 24 éd., Périgueux. Rossi V.G. 1968. L’Atlantide sommersa. Epoca, Sorgial P. 1995. Guia das Ihlas de Cabo Verde. (LXX) 909: 43-62. LIDEL, Editora Técnica Limitada, Lisbon. Rowe N. 1996. The pictorial guide to the liv- Spencer F. 1995. Pithekos to Pithecanthropus: an ing primates. Pogonias Press, East Hampton abbreviated review of changing scientifi c views (NY). on the relationship of the anthropoid apes to Schaller G.B. & Vrba E.S. 1996. Description of Homo. In Corbey R. & eunissen B. (eds): the giant Muntjac (Megamuntjacus vuquangen- Ape, man, apeman: changing views since 1600: sis) in . J. Mammal., 77: 675-683. evaluative proceedings of the symposium, pp. Scortecci G. 1939. Gli iracidi in Libia. Annali del 13-27. Leiden, the , 28 June-1 Museo Libico di Storia Naturale, 1: 382-387. July 1993. Department of Prehistory, Leiden Scortecci G. 1940. Biologia del Tibesti. Annali University, Leiden. del Museo Libico di Storia Naturale, 2: 257- Starck D. & Frick H. 1958. Boebachtungen an 275. aethipoischen Primaten. Zool. Jb. Syst., 85: Sclater P.L. 1858. On the general geographical 41-70. distribution of the members of the class Aves. Sunding P. 1970. Elementer i Kanarioyernes J. Proc. Linn. Soc., 2: 130-145. fl ora, og teorier til fotklaring av fl oraens Sclater P. 1866. Remarks on some monkeys upprinnelse. Blyttia, 28: 229-259. received from St. Kitts, West Indies. Proc. Zool. Sunding P. 1979. Origins of the Maraconesian Soc. London (1866): 79-80. Flora. In Bramwell D. (ed): Plants and islands. Sclater P. 1900. Mr. Sclater on Macacus inuus. Academic Press, London: 13-40. Proc. Zool. Soc. London, 1900: 773-774. Swindler D. 2005. Primate Dentition: An Introduc- Scholfi eld A.F. 1959. Aelian. On the characteris- tion to the Teeth of Non-Human Primates. (Cam- tics of animals, III. William Heinemann Ltd, bridge Studies in Biological and Evolutionary London/Harvard University Press, Cambridge, Anthropology). Cambridge University Press. Massachusetts. Szalay F.S. & Delson E. 1979. Evolutionary his- Segesser F. von, Scheff ran W. & Martin R.D. tory of the primates. Academic Press New York 1995. Parantage analysis within a semi-free- and London. ranging group of Barbary macaques Macaca omas O. 1900. On the mammals obtained in sylvanus. Mol. Ecol., 4: 115-120. South-western Arabia by Messrs. Percival and Segesser F. von, Ménard N., Gaci B. & Martin Dodson. Proc. Zool. Soc. London, VII: 95-97. R.D. 1999. Genetic diff erentiation within nad omas O. 1925. On the mammals (other than between isolated Algerian subpopulations of ruminants) collected by Capt. A. Buchanan barbary macaques Macaca sylvanus: evidence during his second Saharan expedition and pre- from microsatellites. Mol. Ecol, 8: 433-444. sented by him to the National Museum. Ann. Serio-Silva J.C. 2002. Las Islas de los Changos (the Mag. Nat. Hist. London, 9: 187-197. Monkey Islands): the economic impact of eco- Toynbee J.M.C., 1973 – Animals in Roman Life tourism in the region of Los Tuxtlas, Veracruz, and Art. ames and Hudson, London. Mexico. Am. J. Primatol., 68, 5: 499-506. Trevisan Grandi G., Mariotti Lippi M. & Mer- Silsby J. 1980. Inland birds of Saudi Arabia. curi A.M. 1998. Pollen in dung layers from Immel Publishing, London. rockshelters and caves of Wadi Teshuinat (Lib- Smith H.S. 1969. Animal domestication and yan Sahara). In M. Cremaschi & S. Di Ler- animal cult in dynastic Egypt. In Ucko P.J. & nia (eds): Wadi Teshuinat. Palaeoenvironment Dimbleby G.W. (eds): The domestication and and prehistory in south-western Fezzan (Libyan exploitation of plants and animals, 307-314. Sahara), 95-106. C.I.R.S.A. Centro Interuni- Gerald Duckworth & Co Ltd, London. versitario di Ricerca per le Civiltà e l’Ambiente

www.isita-org.com 76 The primates of the western Palaearctic

del Sahara Antico. Edizioni All’Insegna del AETFAT/UNSO Vegetation Map of Africa (3 Giglio, Firenze. C.N.R. Quaderni di Geodi- plates, Northwestern Africa, Northeastern Africa, namica Alpina e Quaternaria, 7. and Southern Africa, 1: 5,000,000). UNESCO, Tutin T.G., Heywood V.H., Burges N.A., Valen- Paris. tine D.H., Walters S.M. & Webb D.A. (eds). Wildman D.E. 1999. Pleistocene dispersal of Afri- 1964. Flora europaea. Cambridge. can hamadryas baboons onto the Arabian penin- Van der Kuyl A.C., Dekker J.T. & Gouldsmit J. sula. Am. J. Phys. Anthropol., Suppl. 28: 277. 1996. St. Kitts green monkeys originate from Willemsen C.A. 2000. L’enigma di Otranto. Il West Africa: genetic evidence from faeces. Am. mosaico pavimentale del presbitero Pantaleone J. Primatol., 40: 361-364. nella cattedrale. Mario Congedo editore, Vaurie C. 1959-1965. e birds of the Palaearctic Galatina (Lecce). fauna. Non –Passeriformes. London. Wilson D.E. & Reeder D.M. (eds). 2005. Mam- Veen M. van der, Grant A. & Barker G. 1996. mal species of the world. A taxonomic reference. Romano-Libyan : Crops and Ani- ird edition. Volume 1. e John Hopkins mals. In G. Barker, D. Gilbertson, B. Jones University Press, Baltimore. & D. Mattingly (eds): Farming the desert. The Yalden D.W., Largen M.J., Kock D. & Hill- UNESCO Libyan Valleys Archaeological Survey. man J.C. 1996. Catalogue of the mammals Volume One, 227-263. UNESCO Publishing, of Ethiopia and Eritrea. 7. Revised Checklist, Paris/Department of Antiquities and Society zoogeography and conservation. Trop. Zool., 1, for Libyan Studies, Tripoli. 9: 73-164. Veracini C. & Masseti M. 2007. Early appearance Zaccagnini C. 1978. Lo scambio dei doni nel of South-American monkeys in Catholic Italy: Vicino Oriente durante i secoli XV-XIII. Roma. the second rediscovery of the Marcgraves’s Zhang Y., Quan G., Lin Y. & Southwick C.H. capuchin monkey, Cebus flavius (Schreber, 1989. Extinction of rhesus monkeys (Macaca 1774). Abstracts of the 12th Brazilian Con- mulatta) in Xinglung, North China. Int. J. Pri- gress of Primatology. Belo Horizonte, 22-23 matol., 10: 375-381. July 2007: 246. Zeuner F.E. 1952. Monkeys in Europe. Past and Waters S.S., Aksissou M., El Harrad A., Hob- present. Oryx, 1: 265-273. belink M.-E. & Fa J.E. 2007. Holding on the Zinner D., Pelaez F., Torkler F. & Berhane D. Djebela: Barbary macaque Macaca sylvanus in 2001. Primates of Eritrea: current distribution northern Morocco. Oryx, 41: 106-108. and habitat. Afr. Primates, 5: 7-17. Wendt H. 1959. L’altra storia della terra. Aldo Martello editore, Milano. White F. 1983. The vegetation of Africa, a descrip- Associate Editor Spartaco Gippoliti & tive memoir to accompany the UNESCO/ Editor, Giovanni Destro Bisol M. Masseti & E. Bruner 77 dated to about 1470-1475 (London, National Gallery). National 1470-1475 (London, about to dated Plate 1 - George Stubbs “Drill and albino hamadryas baboon” (1770-1775). Reproduced by permission of the Royal College of Surgeons of of Surgeons of College Royal the of Kings permission by Reproduced the (1770-1775). of baboon” hamadryas albino and Adoration “Drill Stubbs George - 1 Plate painting “tondo” the of background the in portrayed baboon olive one of Detail frame. small the In London. England, , Botticelli by Sandro

www.isita-org.com 78 The primates of the western Palaearctic (1516-1565). (1516-1565). b) Artistic representation of a Konrad Konrad Gessner Historia Historia animalium by Historia Naturalis Aegypti (1581-1584). Naturalis Alpin in his Historia M. sylvanus , published by Prosper specimen of captive Plate Plate 2 - a) Portrait of a Barbary macaque from the M. Masseti & E. Bruner 79

Plate 3 - Detail of the mosaics of the Great Palace of Istanbul (fi rst half of the 6th century).

www.isita-org.com 80 The primates of the western Palaearctic Jan van der Venationes Venationes Ferarum, Avium, Piscium by the Flemish artist Plate Plate 4 - 16th century engraving representing a monkey hunt from also called Antonio Stradano. Straedt, M. Masseti & E. Bruner 81

Plate 5 - One of the two images of Barbary macaques realized in polychrome marble by the sculptor Cosimo Fancelli around 1555, in the Grotta degli Animaliof the Medici Villa of Castello, near Florence

www.isita-org.com 82 The primates of the western Palaearctic Plate 6 - Detail of the frescoes painted in the fi rst hall of Buontalenti’s Grotta Grandeby Bernardino Poccetti between 1586 and 1587. Boboli 1587. and 1586 between Poccetti Bernardino Grandeby Grotta Buontalenti’s of hall rst fi the in painted frescoes the of Detail - 6 Plate Storici e Artistici Beni i per e Paesaggio del e Architettonici Beni i per Soprintendenza of courtesy Giotti; Lorenzo (photo Florence Gardens, e Prato). di Firenze, Pistoia per le Province e Demoetnologici M. Masseti & E. Bruner 83 Plate 7 - Detail of the wall-paintings of the tomb of Khnemhotep, at Beni Hasan (), showing olive baboons on a fi g tree (Dynasty XII, XII, (Dynasty tree g fi a on baboons olive showing (Egypts), Hasan Beni at Khnemhotep, of tomb the of wall-paintings the of Detail - 7 Plate BC). 1991-1785

www.isita-org.com 84 The primates of the western Palaearctic Plate 8 - Top left: Among the artistic production of the Round Head style, another monkey profi le was initially recognised at Ti-n-ascig, but the the but Ti-n-ascig, at recognised initially was le profi monkey another style, Head Round the of production hunting, artistic monkey red the a of Among left: representation Top alleged - an 8 is there Plate (Libya) Fezzan Farden, In at right: Top carnivore. large a probably more is Late image so-called of the context artistic the with connected been has representation This to assess. cult Pastoral Middle the diffi and to stylised, dating is very image possibly the but III, Teshuinat of site the at drawing wall a in represented baboons four of group a Bottom: style. Pastoral females. juveniles or by three is an adult male, followed rst fi the suggest that (6000-5000 ybp). The mane and sexual characteristics M. Masseti & E. Bruner 85 Plate 9 - The hamadryas evoked in the (Dynasty XVIII). wall-paintings of Tutankhamun’s burial chamber, in the Valley of the Kings at Thebes, Egypt

www.isita-org.com 86 The primates of the western Palaearctic

Plate 10 - The description of Linnaeus and the Latin name of the species, Simia sabaea, (Systema Naturae, 12th ed., 1: 38) were based mainly on a single specimen originating from the island of Santiago, which was published by George Edwards in his Gleanings of Natural history (1758-1764). M. Masseti & E. Bruner 87

Plate 11 - Adult individual of C. sabaeus, depicted by Andrea del Sarto in the Medici Villa of Poggio a Caiano, near Florence (around the end of the second decade of the 16th century).

www.isita-org.com 88 The primates of the western Palaearctic

Plate 12 - Detail of the wall-paintings of the 18th Dynasty tomb of of Rekh-mi-Rē, vizier of Thutmosis III and Amenhotep II (from about 1470 to 1445 B.C.), at Thebes (Egypt). M. Masseti & E. Bruner 89

Plate 13 - Detail of the decoration of the tomb Tiy, the queen of Amenhotep III (1391-1353).

www.isita-org.com 90 The primates of the western Palaearctic

Plate 14 - “Blue monkeys” are a recurrent motif in the production of the Aegean Late Bronze Age artists. M. Masseti & E. Bruner 91 in the work of the 16th century Italian naturalist Ulisse Aldrovandi (V, a., c. 21) - still C. aethiops preserved in the Biblioteca Universitaria of Bologna (Italy) – provides evidence of the conspicuous trade contacts that still existed in the Africa. Egypt and eastern North Middle Ages between Italy, Plate Plate 15. The occurrence of an image of

www.isita-org.com