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2008 ). There is an insular clade, constituting the Indonesian and Phillipine
macaques, and a continental clade, consisting of the macaques from mainland
Southeast Asia.
Ethnophoresy
The close association of long-tailed macaques with humans and their ability to
adapt well to differing habitat types has led to the successful colonization of
several island habitats beyond their natural range due to ethnophoresy, or the
process of an organism being dispersed outside of its range by human trans-
portation (Heinsohn, 2003 ). Ethnophoresy has caused the colonization of long-
tailed macaques on at least five islands in the last several centuries ( Figure
1.1 ) - Mauritius (Sussman and Tattersall, 1981 ; Sussman and Tattersal, 1986 ),
Ngeaur (a.k.a. Anguar) Island in Palau (Poirier and Smith, 1974 ; Wheatley
et al ., 1999), West Papua (Kemp and Burnett, 2003 ; Kemp and Burnett, 2007 ),
Tinjil Island near Java (Kyes, 1993 ), and Kabaena island off Sulawesi (Froelich
et al ., 2003 ). In addition, humans have also carried long-tailed macaques to the
Kowloon Hills of Hong Kong, where they hybridized with rhesus macaques
(Southwick and Southwick, 1983 ; Wong and Ni, 2000 ). These documented
modern cases of colonization provide some evidence that ethnophoresy could
have been the cause of earlier pre-historical colonization events of long-tailed
macaques to islands east of the Wallace line, such as Lombok, Flores, and East
Timur (see Kawamoto et al ., 1984 for discussion).
Republic of Mauritius, East Africa
The first recorded event of long-tailed macaques establishing a satellite popu-
lation on an island outside their range was in Mauritius off the east coast of
Africa (Sussman et al ., Chapter 8). Portuguese traders from Java or Sumatra
probably carried the monkeys during the 1500s along with other forms of exotic
wildlife, such as black rats ( Rattus rattus ) and pigs ( Sus scrofa ) (Sussman and
Tattersall, 1981 ; Tosi and Coke, 2007 ). After this introduction, the macaque
population is believed to have exploded and by the early 1600s M. fascicularis
was well established on the island (Sussman and Tattersall, 1981 ). Although
not documented clearly, the population explosion likely occurred because of
a mixture of human-induced habitat destruction and the availability of abun-
dant food resources from the farms of European settlers. An indication that the
Mauritian macaques largely relied on the settlers of Mauritius is evident by
claims from historians that the early Dutch settlers abandoned Mauritius after
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