JASs Invited Reviews Journal of Anthropological Sciences Vol. 87 (2009), pp. 33-91 The primates 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 Chlorocebus, and possibly also to Erythrocebus and Galago. e data for the present research were collected through a review of all previous knowledge of the primates of this biogeographical region, including their history, and through original sightings and direct observation of fi eld signs. Surveys were carried out directly in North Africa, the peninsula of Gibraltar, and in the Sahara. Additional data on primate 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 human activities and needs. Keywords – Ethnozoology, Macaca, Papio, Chlorocebus. “Hamadryads. A type of wood-nymph. of Eurasia north of the Himalaya along northern All dryads are concerned with trees and live in Africa, including the northernmost part of the them when they are roaming the forest Sahara (cf. Corbet, 1978). Like all faunal regions, - but each hamadryad is the spirit of a however, there can be no precise defi nition of particular tree. 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 (Saudi Arabia), are not com- It can be defi ned approximately as the continent 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 (Algeria), and the Tibesti not extend beyond 30°26’N, 102°52’E (Groves, (Chad). 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 Mauritania, continues across Mali at the same lati- of the Japanese macaque (apart from the “every- tude and thereafter follows the political boundaries where” ape called Homo sapiens). is comprises so including Algeria, Libya and Egypt entirely and most of the Japanese archipelago, including even excluding the whole of Niger, Chad and Sudan. the southern areas of the island of Hokkaido, e entire Arabian peninsula is included, as are located above the 42°N parallel, at the north end the Ahaggar mountains (Algeria), while most of of Japan 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 Yunnan Palaearctic Region, Corbet (1978) includes the (China), north of 25°N, whereas the distribu- archipelagos of Spitzbergen, Iceland, the Azores, tion of the Assam 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 Nepal east ing Vaurie (1959-1965), Cramp (1977) suggests into Tibet 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 macaques and 1 baboon. In fact, for the subcontinent, such as Gujarat and Rajasthan, genus Macaca, he considers the two species M. are not biogeographically comprised within the sylvanus (L., 1758), and M. fuscata, respectively boundary of the Palaearctic, they nevertheless isolated in North Africa 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 Asia, from Afghanistan to Masseti, 2002b); south of the Sahara, the moun- southern China, and probably introduced in an tainous slopes of east Africa 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), Barbary macaque, Macaca sylvanus (L., 1758), is the hamadryas, Papio hamadryas (l., 1758), a the Japanese macaque, Macaca fuscata (Blyth, species still dispersed in eastern Africa and in the 1875), and possibly the snub-nosed monkey, 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 Red Sea, and the Afrotropical e Western Palaearctic is the western por- enclave between Dhofar (Oman) and the Mahral tion of the Palaearctic zoogeographic region Province of Yemen – together with part of Iran (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 Maghreb 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 humans, such the M. sylvanus colony of Gibraltar north-eastern Atlantic Ocean and in Arctic 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 Sahel 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- Ennedi, where the Afrotropical element predomi- tory, through original sightings and direct nates. To the east, the boundary could be limited observation.