AJP 346

American Journal of Primatology 43:181–209 (1997)

RESEARCH ARTICLES
Confirmation and Location of the Hybrid Zone Between
Wild Populations of Macaca tonkeana and Macaca hecki
in Central Sulawesi, Indonesia
E.L. BYNUM,1* D.Z. BYNUM,2 AND J. SUPRIATNA3
1
Department of Anthropology and School of Forestry and Environmental Studies,
Yale University, New Haven, Connecticut
2
Arnold Engineering Development Center, Department of Natural Resources,
Tullahoma, Tennessee
3
Department of Biology, University of Indonesia

Reports of hybridization between Macaca tonkeana and Macaca hecki

were investigated in Central Sulawesi, Indonesia. We defined sets of mor-
phological traits that were diagnostic for M. tonkeana and M. hecki and
then located an areas where animals had intermediate or mosaic fea-
tures. Hybridization as indicated by morphology was detected between
M. tonkeana and M. hecki. The hybrid zone appeared to be strongly cen-
tered at the road that crosses the isthmus of Central Sulawesi from
Tawaeli to Toboli. Macaques in this region were not morphologically uni-
form; animals from the western area of the Tawaeli–Toboli road resembled
M. hecki, while animals from the eastern area resembled M. tonkeana.
The hybrid zone was found to be smaller than previously thought, with
maximum dimensions of approximately 15 and 7.5 km. Clines for diag-
nostic morphological features were broadly coincident, suggesting that
the hybrid zone originated by secondary contact. Analysis of three mu-
seum specimens collected in 1916 provided evidence that the hybrid zone
has been in existence since at least then. The narrow width of the hybrid
zone, along with its age, suggested that some prezygotic or postzygotic
barrier must exist to full introgression between M. tonkeana and M. hecki.
Am. J. Primatol. 43:181–209, 1997. © 1997 Wiley-Liss, Inc.

Key words: Sulawesi macaque, hybridization, Macaca tonkeana, Macaca

hecki

INTRODUCTION
“The macaque monkeys of Sulawesi . . . are remarkable for
the amount of morphological variability they exhibit. These pri-
mates occupy less than 2% of the total area inhabited by the

Contract grant sponsor: National Science Foundation.

*Correspondence to: E.L. Bynum, University of the South, 735 University Avenue, Sewanee, TN 37383.
Email: ebynum@sewanee.edu

Received for publication 4 November 1996; revision accepted 3 July 1997.

© 1997 Wiley-Liss, Inc.

182 / Bynum et al.

macaques as a whole, yet the amount of morphological differen-

tiation they display is easily comparable to that for all non-Su-
lawesi macaques.” (Albrecht 1978)

Only two primate genera, Tarsius and Macaca, are represented on the
island of Sulawesi in Indonesia. Sulawesi is home to at least seven morpho-
logically distinct forms of macaque [Fooden, 1969; Albrecht, 1978] (Fig. 1).
Debate on the taxonomic status of the seven forms has continued due to dif-
ferent interpretations of morphological and genetic variation among them
[Napier & Napier, 1967; Fooden, 1969; Thorington & Groves, 1970; Albrecht,
1977; Groves, 1980].
Recent reports indicate that Sulawesi may be unique in the geographic range
of the genus Macaca. Hybrid or polyspecific groups are rarely seen in undis-
turbed, wild populations of macaques, despite the fact that fertile hybrids have
been produced between many species in captivity [Gray, 1972; Chiarelli, 1973;
Bernstein & Gordon, 1980; Eudey, 1980; Fooden, 1980; reviewed in Fa, 1989]. In
contrast, Groves [1980], Ciani et al. [1989], Watanabe and Matsumura [1991],
Watanabe et al. [1991a, b], and Supriatna and colleagues [Supriatna et al., 1990;
Supriatna, 1991; Froehlich & Supriatna, 1996] have reported that substantial
gene flow and intergradation were taking place between the different forms of
Sulawesi macaque, as evidence by morphologically and genetically intermediate
captive individuals in Sulawesi. The central puzzle of the evolution of Sulawesi
macaques can be stated as the juxtaposition of considerable morphological diver-
sity in the face of the potentially homogenizing influence of hybridization.
Accordingly, in November 1989, an investigation of reported hybridization
between Macaca tonkeana and Macaca hecki was initiated in Central Sulawesi,
Indonesia. Traditionally, hybridization and hybrid zones have been examined with
the aim of delineating species boundaries between related forms or as arenas to
study mechanisms of reproductive isolation and speciation [Arnold, 1992]. Only
more recently have hybrid zones been viewed as “dynamic evolutionary theaters”
[Patton, 1993] and hybridization as a fundamental evolutionary process [Barton
& Hewitt, 1985; Hewitt, 1988; Arnold, 1992]. Although we will not ignore the
taxonomic implications of this work, we are most interested in this latter per-
spective. Thus, we framed several principal questions. Does a hybrid zone exist
between wild populations of M. tonkeana and M. hecki? Does the hybrid zone
have an internal structure, and what pattern of geographic variation is exhibited
by the hybrid zone as a whole? What inferences can be drawn concerning the
origin of the hybrid zone? What evolutionary, historical, environmental, or be-
havioral factors might explain the observed pattern of geographic variation in
the hybrid zone, and, finally, what is the future of the hybrid zone? This paper
will address the first three questions. The last two questions will be addressed in
subsequent publications.

Origin and Differentiation of Sulawesi Macaques

Various scenarios have been proposed for the dispersal and differentiation of
macaques within Sulawesi. Fooden [1969] proposed that Sulawesi was first colo-
nized by a proto–M. tonkeana–like animal from Borneo, while Albrecht’s [1978]
analysis of craniofacial morphology suggested M. maurus as the strongest candi-
date for the basal stock of the Sulawesi macaques. On the basis of variation at
two sites in the beta chains of the hemoglobin molecule, Takenaka et al. [1987a,
b] postulated that M. ochreata and M. hecki were most closely related to M.
 Sulawesi Macaque Hybridization / 183

nemestrina. More characters would be required to clarify relationships among

the other taxa. Fooden and Lanyon [1989] compiled published blood–protein al-
lele frequencies for 17 macaque taxa, including all seven Sulawesi forms. Their
jackknife strict-consensus tree analysis for Sulawesi taxa (22 polymorphic loci,
Macaca nemestrina as an outgroup) identified M. ochreata as the sister taxon of
all other Sulawesi forms. M. maurus and M. tonkeana were found to be closely

Fig. 1. Ranges of Sulawesi macaques (dotted lines) and selected collection locations for Macaca hecki
(Matschie 1901) and Macaca tonkeana (Meyer 1899). Based on Fooden’s [1969] analysis of museum material,
the historical southern collection boundary for M. tonkeana was Uru (3°30´S, 119°53´E), and southeastern
collection boundaries were Palopo (3°01´S, 120°13´E) and Tonkean (1°24´S, 122°30´E). The northern collec-
tion boundary for M. tonkeana, from specimens collected in 1916 by H.C. Raven at “Labua Sore” (correctly
Labuan Sore), was mistakenly reported [Fooden, 1969] as 0°27´S, 120°03´E; the actual location was 0°37´S,
120°03´E. From Fooden [1969], the southern collection boundary for M. hecki was Kampungbaru (1°02´N,
120°48´E), with the eastern collection boundary at Kwandang (0°50´N, 122°53´E). Based on examination of
captive specimens, Groves [1980] extended the known range of M. hecki south to approximately 0°45´S.
Groves also extended the range of M. hecki east past Gorontalo (approximately 123°03´E). Source: Fooden
[1969], Groves [1980], and this study.
184 / Bynum et al.

related to one another, as were M. nigra and M. hecki. However, branching order
remained unresolved in this allozyme analysis.
To date, neither genetic or morphological data have produced consensus on
the sequence or process of differentiation in Sulawesi macaques. A number of
geographical and climatic factors have been discussed in this regard, and all
proposed scenarios have assumed that differentiation of the Sulawesi macaques
proceeded in allopatry. Some authors have hypothesized that many of the cur-
rent lowlands of Sulawesi would have been submerged during warm periods of
the Pleistocene and the highland areas of the current island would have formed
an archipelago [Fooden, 1969]. Circumstantial evidence for the importance of sea
level variation in Sulawesi macaque evolution is provided by marine deposits
and fossils reported at a number of sites above current sea level [Sarasin & Sa-
rasin, 1901; van Bemmelen, 1949].
Some evidence indicates that sea levels around Sulawesi may not have risen
more than 25 m above current levels over the past 2 million years [Whitten et
al., 1987a]. However, some evidence indicates that the extent of uplift on Su-
lawesi during the Quaternary has been greater than that in neighboring areas,
such as Borneo [van Bemmelen, 1949]. In spite of the fact that uplift is an epi-
sodic rather than continuous phenomenon, usually only average rates can be
determined. Tjia et al. [1974] suggested that parts of Sulawesi have uplifted at
the rate of 0.7–9.9 mm per year over the last 11,000–24,000 years. These authors
also proposed an average rate of 4.5 mm/year for eastern Indonesia; over a pe-
riod of 100,000 years, this would have resulted in a positive elevation change in
450 meters.
The Pleistocene archipelago theory will remain incomplete until the role of
uplift in changing land exposure in Sulawesi is more thoroughly understood. Other
authors have noted that warm periods of the Pleistocene were far less common
than cold periods. These researchers have focused on the role of Pleistocene gla-
cial phenomena in creating “islands” of macaque-suitable forest in Sulawesi
[Eudey, 1980; Hamada et al., 1987]. During colder, drier periods, the range of
macaque-suitable forests may have been restricted to the highlands of the inte-
rior and effectively isolated macaque populations. Neither of these theories has
been evaluated with modern tectonic or palynological data.

Variation and Hybridization in Sulawesi Macaques

Fooden [1969] recognized seven species of Sulawesi macaques in his compre-
hensive examination of museum specimens. He found the Sulawesi monkeys to
be strikingly variable in perineal morphology and coloration, in patterns of hair
color in the trunk, forelimb, and crown regions, and in cranial dimensions (Table
I). Albrecht [1977, 1978] documented distinctiveness in craniofacial morphology
among museum specimens of Sulawesi macaque that was a great as the varia-
tion among all non-Sulawesi taxa. Among Sulawesi macaques, craniofacial varia-
tion was “dominated by shape-related differences, with only a minor component
related to size differences” [Albrecht, 1978, p. 81]. Albrecht hypothesized that
cranial variation may be important in species recognition, as the distinct mor-
phologies of the crania of Sulawesi macaques were not related to any known
feeding or ecological differences.
Indeed, the considerable variation in external morphology of Sulawesi
macaques has never been linked to any functional ecological adaptation. As far
as is known, all taxa are similar in body size, dental morphology, and skeletal
adaptations to locomotion. Historical and modern observers have reported all
TABLE I. External Morphological Variation in Museum Specimens of Sulawesi Macaques, after Fooden [1969] and Albrecht [1978]
Taxon Coronal head crest Forearms Shanks Trunk color
M. maurus Absent Brown Brown Brown
M. ochreata Absent Pale grey/brown Pale grey/brown Blackish
M. brunnescens Absent Pale grey/brown Pale grey/brown Brown
M. tonkeana Less well developed Blackish Blackish Blackish
M. hecki Less well developed Dark brown Brown or pale grey/brown Blackish
M. nigrescens Prominent, elongated Brown Brown Brown
M. nigra Prominent, elongated Blackish Blackish Blackish
Taxon Ischial callosities Gluteal fields Rump patch Tail length
M. maurus Suboval, undivided Large Small, not sharply defined Long (42–55 mm)
M. ochreata Suboval, undivided Small Broad, clearly defined Long 35–40 mm)
M. brunnescens Suboval, undivided Small Broad, clearly defined Long (35–55 mm)
M. tonkeana Suboval, undivided Small Broad, clearly defined Long (40–70 mm)
M. hecki Reniform, partially subdivided Rudimentary to absent Weakly defined, dorsally obsolete Short (13–36 mm)
M. nigrescens Suboval, undivided Rudimentary to absent Absent Short (25 mm)
M. nigra Reniform, completely divided Rudimentary to absent Absent Short (20 mm)
Sulawesi Macaque Hybridization / 185
186 / Bynum et al.

Sulawesi taxa as primarily forest-dwelling creatures, although some taxa are

also known to raid crops and have been sighted in other habitats such as beaches
and grasslands–scrubland [Fooden, 1969; Groves, 1980; Watanabe & Brotoisworo,
1982; Supriatna, 1991]. All Sulawesi macaques studied are highly frugivorous,
although leaves, flowers, invertebrates, and possibly small vertebrates are also
eaten [Muskita, 1988; Zahrah, 1988; Hadi, 1989; Supriyadi, 1989; Supriatna, 1991;
Kohlhaas, 1993].
Behaviorally, Sulawesi macaques share a number of highly developed and
unusual facial expressions and patterns of affiliative embracing and reconcilia-
tion [Dixson, 1977; Hadidian, 1980; Thierry, 1984, 1985a, 1986; Baker & Estep,
1985; Herrenschmidt, 1985; Baker & Bynum, 1989; Petit & Thierry, 1992; Petit
& Thierry, 1994a, b]. Sexual patterns appear to be broadly similar in Sulawesi
macaque taxa studied to date [Dixson, 1977; Thierry, 1986; Baker & Bynum,
1989; Thierry et al. 1994]. With the possible exception of the loud call or chirp
bark that appear to be different in each taxon [Watanabe & Brotoisworo, 1982;
personal observation], no other behavior patterns are known to be taxon-spe-
cific. Captive hybrid monkeys appear to share behavioral traits with each of
their parental taxa, but the relative frequency of expression of elements of
the agonistic, affiliative, and sexual repertoire in hybrids is not known [Baker
& Bynum, 1989].
Groves [1980] was among the first to attempt field work on Sulawesi macaques,
with emphasis on the borderlands between taxa. In most presumed border areas,
observation of wild populations proved unfeasible or unsatisfactory. Accordingly,
Groves focused on assessing pet monkeys by external morphological criteria adapted
from Fooden [1969]. Groves observed no morphological intergradation in wild or
captive M. maurus or M. tonkeana but did not observe captive animals that ap-
peared intermediate in morphology at the borderlands between M. tonkeana and M.
hecki and M. nigra and M. nigrescens. Groves therefore recognized four species of
Sulawesi macaque: M. maurus, M. tonkeana (with M. t. tonkeana and M. t. hecki as
subspecies), M. nigra (with M. n. nigra and M. n. nigrescens as subspecies, and M.
ochreata (with M. o. ochreata and M. o. brunnescens as subspecies). Groves’ study
provided the first evidence for in situ hybridization between some of the Sulawesi
macaque taxa, although the short period of research and the small samples sizes
observed in each area precluded definitive conclusions.
Ciani et al. [1989] surveyed five Sulawesi taxa (M. maurus, M. tonkeana, M.
hecki, M. nigrescens, and M. nigra) and several borderlands between forms in
1983. Data were collected for both morphological and genetic traits in a number
of captive individuals which were typically reported by the owners as caught in
nearby forests. Results suggested restricted regions of steep clinal change in mor-
phological score (areas of intergradation) bounded by areas of little variation
(“pure” forms). These data suggested that gene flow was occurring to at least
some extent between the forms investigated, including M. tonkeana and M. mau-
rus, which Groves (1980) found not to intergrade.
Watanabe and colleagues [Watanabe & Matsumura, 1991; Watanabe et al.,
1991a, b] have surveyed extensively in Sulawesi since 1980, examining pets and
observing wild populations where possible in an effort to locate borderlands be-
tween Sulawesi taxa. Watanabe et al. [1991a] provided convincing evidence for
phenotypic intergradation between M. tonkeana and M. hecki based on examina-
tion of captive specimens. Their examination of the M. tonkeana/M. ochreata bor-
der zone (1991b) was less systematic and involved smaller sample sizes than the
M. tonkeana/M. hecki study but provided preliminary evidence of some degree of
morphological intergradation within the zone.
 Sulawesi Macaque Hybridization / 187

Supriatna and colleagues [Supriatna et al., 1990; Supriatna, 1991; Froehlich

& Supriatna, 1996] have conducted the most detailed investigation of geographic
variation and intergradation in Sulawesi macaques to date. Research at Maiwa
in South Sulawesi revealed phenotypically mixed troops, mixed bachelor bands,
and mating between animals of M. tonkeana and M. maurus phenotype.
Supriatna and colleagues [Supriatna et al., 1990; Supriatna, 1991; Froehlich
& Supriatna, 1996] extended their study to include samples from populations of
M. tonkeana, M. maurus, and their putative hybrids along a 400 km transect.
Hybrid animals were found in the area between the distributions of the parental
populations, in an area at least 10 km from north to south in the western area of
the southern peninsula, and at least 35 km north to south in the eastern area of
the southern peninsula.

Macaca tonkeana and Macaca hecki: Summary of Previous Research

Selected historical collection locations and currently understood range bound-
aries for M. tonkeana and M. hecki are presented in Figure 1. M. hecki resembles
M. tonkeana in having “bushy cheek whiskers, a short suberect crest on the crown,
long but broad muzzle, and very great upper facial breadth” [Groves, 1980, p.
105]. Strongly developed supraorbital tori and weakly to moderately developed
supramaxillary ridges also unite M. hecki to M. tonkeana and the other southern
Sulawesi species [Fooden, 1969]. Albrecht [1977] found M. tonkeana and M. hecki
similar in overall skull form. As in other Sulawesi macaques, there is no indica-
tion of gross musculoskeletal or dental differences between the two taxa. Scant
data available on comparative body sizes indicated that head and body lengths of
adults (segregated by sex) averaged only 5% larger in M. tonkeana than in M.
hecki [Fooden, 1969, 1980]. The degree of sexual dimorphism was similar in each
form [Fooden, 1969].
In spite of these similarities, M. hecki and M. tonkeana are phenotypically
distinct. In a number of characters (shape of the baculum, tail vestige as a pro-
portion of body size, ischial callosity shape, ischium form, relative reduction of
the rump patch and gluteal fields), M. hecki resembles the northern species M.
nigra and M. nigrescens more closely than the southern and central species M.
maurus, M. tonkeana, M. ochreata, and M. brunnescens [Fooden, 1969; Groves,
1980]. Electrophoretic and sequence analysis of blood proteins also indicated a closer
affinity between M. hecki and the northern taxa than between M. hecki and the
southern and central forms {Kawamoto et al., 1985; Takenaka et al., 1987a].
Extensive work has been done on the behavior of captive M. tonkeana groups
in Europe and the United States [Herrenschmidt, 1977, 1985; Bayart, 1982; Bayart
& Anderson, 1985; Thierry, 1984, 1985a, 1985b, 1986; Thierry et al., 1989, 1990;
Petit et al., 1992; Masataka & Thierry, 1993; Petit & Thierry, 1994a]. No compa-
rable studies exist for M. hecki. Some authors, including Supriatna [1991], have
stated or implied that M. hecki inhabits and is adapted to drier areas than M.
tonkeana. But information on ecology from wild populations is still fragmentary.
Supriatna [1991] collected population and ecological data on M. tonkeana living
in Lore Lindu National Park in Central Sulawesi as a component of his disserta-
tion research. Watanabe [unpublished manuscript] observed the ecology and be-
havior of unhabituated M. hecki for 8 months in the proximal area of the northern
arm of the island.
When Groves [1980] surveyed the isthmus of Central Sulawesi for the range
boundary between M. hecki and M. tonkeana, he observed no mixed or morpho-
logically intermediate free-ranging groups. However, 16 pet monkeys exhibited
188 / Bynum et al.

various combinations of parental morphological traits and convinced Groves that

intergradation was taking place. Ciani et al. [1989] presented evidence that cap-
tive animals originating in the isthmus of Central Sulawesi had morphology and
gene frequencies for polymorphic loci intermediate to those of M. tonkeana and M.
hecki. A limitation of this data set was the small number of animals examined for
morphological and genetic traits. In addition, intermediate allelic frequencies on the
M. tonkeana/M. hecki border were demonstrated at three loci (PGM-2, GC, HB) and
not at six other variable loci (TF, CA-1, PGM-1, PI, 6-PGD, CA-2).
Watanabe et al. [1991a] evaluated the morphology of a number of pet M.
tonkeana and M. hecki from Central Sulawesi using methods almost identical to
those of Groves [1980]. Watanabe et al. examined over 100 captive animals, some
reported as originating far from the same reported as originating near the puta-
tive border zone. They described several morphologically intermediate captive
animals, all referable to a relatively small area in the isthmus that connects
Central and North Sulawesi.
This portion of the isthmus is characterized by a series of narrow ridges and
valleys oriented east–west. There are no obvious geographical barriers such as
high mountains or large rivers that would serve to isolate macaque populations,
except for a paved road which crosses the isthmus from Tawaeli to Toboli (Fig.
2). Partial conversion of remaining forest to agriculture extends a maximum of
approximately 1–2 kilometers in either direction from the road. The implications
of the road for contact between M. tonkeana and M. hecki were unclear as re-
search began, although from sightings of macaques crossing the road it was clear
that it and associated disturbance did not represent an absolute barrier to
macaque movement.
In summary, as this project began, the relationship between M. tonkeana
and M. hecki was characterized by unexplained morphological and genetic affini-
ties and suspected morphological intergradation. The status of wild populations,
however, had scarcely been examined.

METHODS
Definition of Terms
We use the term hybridization to mean “the interbreeding of [individuals
from] two populations, or groups of populations, which are distinguishable on
the basis f one or more [heritable] characters” [Harrison, 1990, as modified from
Woodruff, 1973]. A hybrid zone is thus an “area of interaction between distinct
groups of individuals, in which these interactions result in at least some off-
spring of mixed ancestry” [Harrison, 1990]. Hybrid zones are made up of one or

Fig. 2. Study locations in Central Sulawesi. At left, regions 1a (Tambu) and 1b (Gunung Bosa) were base-
line M. hecki. Regions 6a (Kamarora) and 6b (Sidaunta) were baseline M. tonkeana. In regions 2 (Sinio), 3
(Avalua), 4 (Tawaeli–Toboli road [TATO]), and 5 (Pangi-Binangga), monkeys were assigned to one of the
parental taxa or identified as hybrids based on diagnostic morphological features. Shaded area indicate
remaining forest cover circa 1985. On the right, kilometers 30–60 of the Tawaeli–Toboli road is shown, and
Puncak Beringin research station at kilometer 56 is indicated. The large rectangle is a maximum estimate
for the length (15 km) and width (7.5 km) of the hybrid zone, based on 1) the forested area of the isthmus in
the area of the hybrid zone and 2) the location of phenotypically parental populations at Avalua and Pangi-
Binangga, respectively. The narrow hatched rectangle at the center is the postulated zone of maximum
morphological variation within the hybrid zone. See text for discussion of the oblique orientation of the
hybrid zone. Compiled from 1) satellite data, radar, airphotographs, and topographic maps by the Depart-
ment of Transmigration, government of Indonesia, 2) map of the Gunung Tanggunung Protected Forest
Boundaries, Directorate General of Forest Inventory and Use, Ministry of Forestry, government of Indone-
sia, and 3) a hand-drawn map of the Pangi-Binangga area, 1984 report of a team from Directorate of Na-
ture Protection and Forest Conservation (PHPA), Palu, Central Sulawesi, Indonesia.
Sulawesi Macaque Hybridization / 189

Figure 2.
190 / Bynum et al.

more clines, with a cline defined as “a geographic gradient in a measurable char-

acter, or gradient in gene, genotype, or phenotype frequency” [Endler, 1977]. As
Harrison [1990] has stated, this working definition makes a minimum of assump-
tions about the history of the interaction and the evolutionary forces now acting
in a given hybrid zone. This will result in the grouping together of superficially
similar phenomena as a starting point for hybrid zone analysis [Harrison, 1990].
A number of different types of characters can be used to define a hybrid zone.
However, in this study, consideration was restricted to nonmetric external morphol-
ogy. Thus, in this study, a hybrid zone would be the area in which at least some
morphological phenotypes were intermediate to or mosaics of parental types, where
a mosaic individual would exhibit “pure” features of both parental taxa.

Sampling Methods
Research began by sampling the morphology of Macaca tonkeana and Macaca
hecki far from the suspected contact zone in order to quantify the extent of in-
trataxon morphological variation before attempting to evaluate animals in the
reported contact area. Tambu and Gunung Bosa (Fig. 2, regions 1a and 1b) were
chosen as baseline populations of M. hecki, and Kamarora and Sidaunta were
chosen as baseline populations for the morphology of M. tonkeana (Fig. 2, re-
gions 6a and 6b).
Methods for locating monkeys and diagnosing morphology were similar in
all surveyed locations. Study areas were searched until contact was established
with a macaque party or individual. Contact was then maintained for as long as
possible. One morphology record was collected for each monkey observed.
Monkeys were evaluated for nonmetric morphological features using criteria
adapted from Fooden [1969] and Groves [1980] (Table II; Fig. 3). These data
were then examined for traits that could be used to consistently differentiate the
taxa. Procedures for identifying diagnostic traits are described in the next sec-
tion. A value of 1 was assigned to the M. hecki condition for each determined
diagnostic trait, a value of 3 was assigned to the M. tonkeana condition, and a
value of 2 was assigned for intermediate features. Having established a suite of
diagnostic traits, we traveled to other areas in Central Sulawesi to examine mor-
phology, taking care to sample on both sides of the suspected hybrid zone.
Data on color state variation in morphological traits were also recorded but were
not used for diagnostic purposes, as field tests indicated high intra- and interobserver
variability in scoring. Individual recognition and reproductive traits such as scars,
patterns of depigmentation on the face, male scrotal color, and female sexual swell-
ing were also recorded in order to avoid repeat descriptions of the same monkey.
Age and sex of contacted individuals were determined at the time of sight-
ing. We chose to focus on describing adults at the expense of other age classes, as
research has suggested that some of the diagnostic features of Sulawesi macaques
may be fully developed only in adults [Hadidian, 1980]. In addition, adults are
larger and therefore have a larger surface area of diagnosable features.
Summary statistics (number of monkeys present and age and sex distribu-
tion) were gathered for each contact. Obtaining accurate counts was difficult for
the largely unhabituated and forest-dwelling monkeys of this study, and in most
cases the relationship between encountered monkeys and the social unit was
unclear. Therefore, we referred to party size for encountered monkeys rather
than group size [cf. van Schaik et al., 1983].
During one phase of the research, we spent 13 months in the Puncak Berin-
gin Study Site (Fig. 2), adjacent to kilometer 56 along the road from Tawaeli to
TABLE II. Morphological Traits Examined for Diagnostic Value*

Trait name
Crown hair orientation
Cheek whisker contrast
Throat contrast
Venter contrast
Forearm contrast
Gluteal fields extent
Ischial callosity shape
Rump patch extent
Shanks contrast
Character states
Figure 3 M. hecki Intermediate M. tonkeana
1 Crown hairs smooth Longer hairs, no distinct crest Distinct crest
2 No contrast to face Contrasting tones Distinct contrast
3 No contrast to surrounding hair Contrasting tones Distinct contrast
4 Distinct contrast to back Contrasting tones No contrast
5 Distinct contrast to upper arm Contrasting tones No contrast
6 Absent Less then 25% size of ischial callosities More than 25% size of ischial
callosities
7 Reniform Distorted oval Oval
8 Interior to midline of thigh Past midline of thigh but not extending Extends to or past sides of
to or past sides thigh
9 Distinct contrast to upper leg Contrasting tones No contrast

*Diagnostic traits are illustrated in Fig. 3. Traits were initially selected based on Fooden [1969] and Groves [1980] and modified for field observations.
Sulawesi Macaque Hybridization / 191
192 / Bynum et al.

Fig. 3. Morphological features of adult Macaca hecki and Macaca tonkeana, with numbered features and
character states as indicated in Table 2. In M. hecki, gluteal fields extent could be either absent [6(1)] or
less than 25% of ischial callosities [6(2)]. Similarly, rump patch extent could be either interior to midline of
thigh [8(1)], or past midline of thigh but not extend to or past sides [8(2)]. Drawing by Karen Petras.
 Sulawesi Macaque Hybridization / 193

Toboli (hereafter TATO). Possible multiple observations of the same monkey were
eliminated as far as possible by identifying individuals by idiosyncratic marks
and features.
In addition, J.W. Froehlich of the University of New Mexico evaluated three
juvenile Sulawesi monkey specimens housed at the Smithsonian Museum using
a slightly modified set of diagnostic traits [Bynum et al., in press]. These speci-
mens were collected in 1916 by H.C. Raven for the United States National Mu-
seum (now the Smithsonian) in “Labuan Sore” on the east coast of the isthmus
that connects Central and North Sulawesi.

Data Preparation and Analysis

A trait was considered diagnostic if no more than 5% of adult M. tonkeana
shared the same character state for this trait with adult M. hecki. In order to
determine whether diagnosis should be restricted to same-age and same-sex
classes across regions, traits were evaluated for a sex-associated or developmen-
tal component using Chi-square tests for independence [Fienberg, 1980; Siegel &
Castellan, 1988].
Counts of character states for diagnostic features in adults were tabulated across
sampled regions and converted to proportions of observations for each feature in
order to equalize sample sizes for comparison. Since individual morphological traits
may or may not have sorted independently, analysis by multivariate comparison of
tabulated character states for all traits by region was selected rather than univariate
comparison of character state data among geographic areas.
Multidimensional scaling (MDS) [Wilkinson, 1990; Morrison et al., 1992] was
used to compare character state counts among regions. Counts and proportions
of character states for each variable were used to generate Pearson product–
moment correlation matrices comparing regions [Sneath & Sokal, 1973]. Corre-
lation matrices were then used to generate MDS graphical output.
The morphology of individual monkeys was compared using a Mean Hybrid-
ity Index (MHI) [Phillips-Conroy & Jolly, 1986; van Zyll de Jong et al., 1995].
The MHI was constructed by averaging the morphological scores of individual
monkeys with four or more of the six diagnostic traits scored. Kruskal-Wallis
one-way analysis of variance [Siegel & Castellan, 1988; Wilkinson, 1990] was
used to compare population means among the six geographic areas sampled. Dif-
ferences in central tendency of population MHI values between pairs of sampled
locations were examined using Wilcoxon-Mann-Whitney tests [Siegel & Castellan,
1988; Wilkinson, 1990].
Graphical and statistical analyses were carried out using Microsoft Excel
4.0, Systat 5.1 [Wilkinson, 1990], and BIOM [Rohlf, 1992].

RESULTS
Sightings and Description of Surveyed Locations
Over the course of the study, there were 627 sightings of wild monkeys (Table
III), in which 4,034 individuals were counted and 2,308 descriptions collected.
Total contact time was over 19,000 min, or about 320 h. Average contact duration
was 31 min, and average party size was 7.15 individuals.
Table IV presents descriptive information (dates surveyed, broadly defined
vegetation types, elevation, topography, and sighting information) for each geo-
graphic area visited. Detailed descriptions of surveyed areas can be found in
Bynum [1995].
194 / Bynum et al.
TABLE III. Summary of Wild Sightings Across All Geographic Areas Surveyed

Total Mean Mean

Number of contact contact Number of party Number of
Location sightings time (min) time (min) individuals size descriptions
Gunung Bosa 1 78 78 6 6.00 4
Tambu 55 1,823 34 417 7.60 214
Sinio 6 202 34 61 10.00 27
Avalua 12 270 25 74 6.20 59
Tawaeli-Toboli 507 15,436 31 3,165 6.40 1,775
Pangi-Binangga 27 589 23 135 5.00 70
Sidaunta 17 759 45 164 10.00 151
Kamarora 2 30 15 12 6.00 8
Total, all locations 627 19,187 35 4,034 7.15 2,308

Diagnostic Traits in Baseline Populations of M. tonkeana and M. hecki

Table V presents character states that were evaluated for adult M. tonkeana
and M. hecki. The variables of Cheek Whisker Contrast, Gluteal Fields Extent,
Ischial Callosity Shape, Rump Patch Extent, and Shank Contrast were diagnos-
tic features according to criteria previously discussed. Crown Hair Orientation
was included as a diagnostic feature despite the greater than 5% overlap be-
tween adult M. tonkeana and M. hecki for character state 1 (crown hairs smooth).
Observations early in the study of two adult female M. tonkeana directly after a
rainstorm accounted for this overlap; later experience suggested to us that the
smooth-crowned appearance of these two individuals was a result of weather
conditions rather than deviation from the typical M. tonkeana phenotype.
No statistically significant differences were found between adult males and
females for any diagnostic feature in M. tonkeana and M. hecki (Chi-square test
for independence with alpha = 0.05; critical value was 5.991 with 2 d.f.). Sample
sizes for comparisons ranged from 19–68 (Table VI). One diagnostic feature for
the M. tonkeana baseline population was found to have a developmental compo-
nent, although the small sample sizes in the nonadult age classes made statisti-
cal comparison problematic (Table VI). Frequencies of character states for Cheek
Whisker Contrast were significantly different between adults and nonadults. Ju-
veniles were more likely than other age classes to have dark cheek whiskers (N
= 85, Chi-square = 26.26, d.f. = 4, P < 0.001). However, this must be interpreted
cautiously due to the small number of juveniles sampled.
No statistically significant age-related differences were noted in M. tonkeana
baseline populations for the feature rump patch extent. However, scoring for this
feature evolved over the initial months of the study. Before March 1990, Rump
Patch Extent was scored as a binary variable rather than a three character state
variable; not all of these earlier data could be directly transferred to the three
character state system. In the initial period, observations were made on four
juvenile individuals whose rump patch differed from the typical M. tonkeana
condition of bushy out to or past the sides of the leg. Therefore, we strongly
suspected that Rump Patch Extent had a developmental component and excluded
it when making comparisons across age classes.
Developmental differences in diagnostic features indicated that caution should
be exercised in applying diagnostic features across age classes. In contrast, no
evidence was found to suggest that diagnostic traits cannot be compared between
adult males and females.
TABLE IV. Study Locations*
Region Location Dates surveyed Elevation range Vegetation Comments
1a Tambu 24 Nov.–01 Dec. 1989 100–500 Lowland forest, mixed forest/ Severe disturbance in some areas
10–14 Dec. 1989 garden, clove and chocolate adjacent to road
22–27 June 1991 monoculture
1b Gunung Bosa 23 Nov. 1989 200–300 Logged lowland forest Moderate to severe disturbance
2 Sinio 02–08 Feb. 1990 300–900 Lowland forest to transition Moderate disturbance at lower
lower montane forest elevations; only slight
disturbance at higher elevations
3 Avalua 13 March 1990 100–600 Lowland forest, grassland on Moderate to severe disturbance at
19–28 March 1990 exposed slopes and all elevations
ridgetops near the coast
4 Tawaeli–Toboli Multiple trips, Dec. 140–1,130 Lowland forest to lower montane Slight to severe disturbance along
1989–May 1990 forest, mixed forest/garden, road and up to a kilometer north
Oct. 1990–Dec. 1991 clove monoculture and south
5 Pangi-Binangga 16–22 Jan. 1990 100–800 Lowland forest Slight to moderate disturbance
6–13 March 1990 at all elevations
Several short trips, 1991
6a Kamarora 12–15 Nov. 1989 Lowland forest, mixed forest/garden
6b Sidaunta 18–21 Nov. 1989 550–950 Lowland forest Slight disturbance, surveyed areas
16–19 Dec. 1989 inside Lore Lindu National Park
09–14 Sept. 1991

*Region numbers refer to Fig. 2. Elevations in meters above sea level.

Sulawesi Macaque Hybridization / 195
196 / Bynum et al.
TABLE V. Proportions of Morphological Character State Variables that were Examined
for Diagnostic Ability in Adults from Baseline Populations of Macaca hecki and Macaca
tonkeana*
Character states
Variable 1 2 3 N
M. tonkeana Crown hair orientation 7 7 86 42
Cheek whisker contrast 0 0 100 69
Throat contrast 0 0 100 32
Venter contrast 14 5 81 21
Forearms contrast 0 0 100 60
Gluteal fields extent 0 0 100 43
Ischial callosity shape 0 0 100 20
Rump patch extent 0 0 100 56
Shanks contrast 0 0 100 54
M. hecki Crown hair orientation 98 2 0 86
Cheek whisker contrast 100 0 0 81
Throat contrast 67 0 33 3
Ventrum contrast 97 0 3 30
Forearms contrast 20 63 17 30
Gluteal fields extent 93 7 0 29
Ischial callosity shape 100 0 0 42
Rump patch extent 30 70 0 23
Shanks contrast 97 3 0 63

*N is number of observations.

Morphology of Baseline Populations and Survey Locations Compared

Figure 4 (top) presents an MDS ordination generated from a Pearson prod-
uct–moment correlation of proportions of character states for adults among
sampled locations 1–6 (Table VII). The MDS generated three clusters of points.
To the left, baseline M. hecki (1) clustered closely with survey populations at
Sinio (2) and Avalua (3). On the right, morphology data from Pangi-Binangga
clustered closely with baseline M. tonkeana. Intermediate to these clusters were
data from the Tawaeli–Toboli road (Fig. 4, 4). In addition, sampled locations dif-
fer from one another primarily along dimension 1 (Fig. 4), which we interpreted
to correspond to a north–south geographic axis.
Average MHI scores increased from north to south (Table VIII). Region 3
(Avalua) and region 5 (Pangi-Binangga) were less than 5 km on either each side
of region 4 (TATO road) (Fig. 4), yet there was a sharp step in average scores
moving across these areas. The variance in the MHI also increased sharply mov-
ing into the TATO region from either direction.
Various transformations of MHI scores across the six areas did not produce
distributions suitable for ANOVA. Therefore, nonparametric Kruskal-Wallis tests
were used to compare MHI across regions. Statistical results confirmed that among
the six geographic regions surveyed, between-location variation in MHI over-
whelmed within-location variation (Kruskal-Wallis test statistic = 161.65, P <
0.001 with 5 d.f.).

Variation in Morphology Along the TATO Road

Morphology in the TATO region was intermediate to that of baseline M.
tonkeana and M. hecki. However, a scatterplot of MHI against corresponding
TABLE VI. Sex and Age Differences in Diagnostic Features in Baseline Populations*
Taxon Character
M. hecki Crown hair orientation
Cheek whiskers contrast
Gluteal fields extent
Ischial callosity shape
Rump patch extent
Shanks contrast
M. tonkeana Crown hair orientation
Cheek whiskers contrast
Gluteal fields extent
Ischial callosity shape
Rump patch extent
Shanks contrast
*Differences were examined using Chi-square tests for independence with alpha = 0.05. Significant nonindependence indicated by box. For 2 × 3 tables, critical value of the
Chi-square was 5.991 (2 d.f.); for 3 × 3 tables, critical value was 9.488 (4 d.f.). See discussion of potential age differences in Rump Patch Extent in text.
By sex By age class
Character state Adult female Adult male N All adult All juvenile All subadult N
1 29 37 84 11 12
2 1 1 2 0 0
3 0 0 68 0 0 0 109
1 25 35 81 13 18
2 0 0 0 0
3 0 0 60 0 0 0 112
1 14 16 27 1 1
2 0 0 2 0 0
3 0 0 30 0 0 0 31
1 14 16 42 5 7
2 0 0 0 0 0
3 0 0 30 0 0 0 54
1 3 3 7 0 1
2 1 9 16 0 1
3 0 0 16 0 0 0 25
1 19 21 61 10 12
2 1 1 2 1 2
3 0 0 42 0 0 0 88
1 2 1 3 0 0
2 1 1 3 0 0
3 17 16 38 36 2 3 47
1 0 0 0 3 0
2 0 0 0 0 0
3 31 25 56 69 6 7 85
1 0 0 0 0 0
2 0 0 0 0 0
3 28 13 41 43 3 1 47
1 0 0 0 0 0
2 0 0 0 0 0
3 9 11 20 20 1 2 23
1 0 0 0 0 0
2 0 0 0 0 0
3 27 23 50 56 5 6 67
1 0 0 0 0 0
2 0 0 0 0 0
3 26 21 47 54 10 6 70
Sulawesi Macaque Hybridization / 197
198 / Bynum et al.

Fig. 4. Top: Multidimensional scaling (MDS) of diagnostic morphology generated from Person product–
moment correlation matrices of proportions of character states among adults across survey regions 1–6.
Bottom: MDS of diagnostic morphology by kilometer along the TATO road. Morphology of baseline M. tonkeana
and M. hecki and regions 2, 3, and 5 provided for context; TT34 is kilometers 30–50, TT50 is kilometers 50–
59, and TT59 is kilometers 59 and above along the TATO road. Locations as indicated in Fig. 2.
TABLE VII. Proportions of Character States Across Sample Regions 1–6 and by Kilometer Along the Tawaeli–Toboli Road (Fig. 2)*
Survey region Tawaeli–Toboli road
Character state 1 2 3 4 5 6 Kilometers 30–50 Kilometers 50–59 Kilometers 59+
Crown hair orientation 1 98 100 85 49 6 7 81 50 23
Crown hair orientation 2 2 0 15 46 11 7 19 46 60
Crown hair orientation 3 0 0 0 5 83 86 0 3 16
N 86 14 26 334 18 42 27 264 43
Cheek whisker contrast 1 100 100 78 45 11 0 93 45 11
Cheek whisker contrast 2 0 0 22 36 26 0 7 37 53
Cheek whisker contrast 3 0 0 0 19 63 100 0 19 37
N 81 14 23 306 27 69 27 241 38
Gluteal fields extent 1 93 100 7 12 0 0 60 11 0
Gluteal fields extent 2 7 0 47 10 0 0 20 8 18
Gluteal fields extent 3 0 0 47 79 100 100 20 81 82
N 29 8 15 136 16 43 5 114 17
Ischial callosity shape 1 100 100 100 24 0 0 80 24 7
Ischial callosity shape 2 0 0 0 9 40 0 0 12 0
Ischial callosity shape 3 0 0 0 67 60 100 20 64 93
N 42 7 18 96 10 20 5 76 15
Rump patch extent 1 30 . 59 10 2 0 20 11 0
Rump patch extent 2 70 . 41 57 0 0 60 52 79
Rump patch extent 3 0 . 0 33 98 100 20 37 21
N 23 0 17 204 44 56 10 161 33
Shanks contrast 1 97 100 77 21 0 0 75 17 5
Shanks contrast 2 3 0 18 16 5 0 19 17 9
Shanks contrast 3 0 0 5 64 95 100 6 66 86
N 63 13 22 200 21 54 16 162 22

*Diagnostic traits are divided into character states as indicated in Table 2 (e.g., Shanks Contrast 2 corresponds to the second of three character states possible for the
feature). Data presented for adults only. N is number of observations. No animals were scored in Sinio (region 2) for Rump Patch Extent.
Sulawesi Macaque Hybridization / 199
200 / Bynum et al.
TABLE VIII. Population Means and Standard Deviations for the Mean Hybridity
Index (MHI)*
Sampled population Region Number MHI S.D.
M. hecki 1 34 1.09 ± 0.01
Sinio 2 7 1.00 ± 0.00
Avalua 3 19 1.26 ± 0.06
Tawaeli–Toboli road (all sections) 4 112 2.15 ± 0.30
Tawaeli–Toboli road kilometers 30–50 4 8 1.25 ± 0.00
Tawaeli–Toboli road kilometers 50–59 4 89 2.17 ± 0.00
Tawaeli–Toboli road kilometers 59+ 4 15 2.54 ± 0.00
Pangi-Binangga 5 12 2.83 ± 0.07
M. tonkeana 6 39 2.96 ± 0.01
*Sampled locations indicated in Fig. 2.

kilometer markers along the TATO road (data were not taken directly along the
road but typically in forests adjacent to kilometer markers) revealed that mor-
phology was not uniform within this region (Fig. 5). A weak positive relationship
existed between location and MHI (N = 112, r = 0.52, P < 0.001). This relation-
ship appeared strongest in the far left and right areas of the plot. Animals in
kilometers 30–40 were more M. hecki–like, and those from kilometers 59 and
above were M. tonkeana–like.
An examination of population MHI by kilometers along the TATO road con-
firmed this pattern (Table VIII). Population MHI differed from each other in the
expected directions, and variance in MHI was highest in the kilometers 50–59

Fig. 5. MHI for adults by kilometer along the TATO road. Kilometer markers as indicated in Fig. 2.
 Sulawesi Macaque Hybridization / 201

region. Kruskal-Wallis one-way analysis of variance confirmed that population

MHI was different (Kruskal-Wallis test statistic = 23.94, P < 0.991 with 2 d.f.).
Without more sample points, in particular near kilometers 40–50, it was dif-
ficult to evaluate whether the extreme variation in MHI in kilometers 50–59 was
a real phenomenon or an artifact of more intensive sampling compared to other
areas. Few sample points were collected for kilometers 40–50 along the TATO
road, as human populations along the road were clustered in this heavily dis-
turbed area.
A population-level comparison of MHI was comprised of data only from mon-
keys scored for four or more of the six possible diagnostic features. In contrast,
multidimensional scaling analysis used counts of character states for all animals
observed. Thus, larger sample sizes were available for MDS analysis, particu-
larly for undersampled areas. MDS analysis confirmed that morphology was not
uniform along the TATO road (Fig. 4, bottom). Monkeys in kilometers 30–50 clus-
tered tightly with M. hecki morphotypes, and animals in kilometers 59 and above
were closer in morphology to M. tonkeana than animals in kilometers 50–59.

Dimensions and Orientation of the Hybrid Zone

A maximum of 15 km for the length of the morphologically detectable hybrid
zone was estimated from the extent of remaining forest cover in the area of the
hybrid zone. A maximum of 7.5 km for width of the morphologically detectable
hybrid zone was estimated from the distance between the parental-type samples
at Avalua (region 3) and Pangi-Binangga (region 5). Observed morphological varia-
tion in the TATO region suggested a hybrid zone with a central zone of high
morphological variation, bounded by areas more like parental forms, and the
oblique orientation indicated in Figure 2. This oblique orientation, although not
the only possible configuration, is also consistent with the east–west constriction of
remaining forest cover in the area north and south of the TATO road.
The regular and rectangular shape of the hybrid zone presented here is use-
ful as a schematic but is almost certainly not a complete description of a zone
likely influenced by fine-scale variation in topography and degree of disturbance.
However, data from this study did not provide sufficient resolution to evaluate
other possibilities.

Morphology and Placement of “Labuan Sore” Specimens

Our identification of M. hecki at the Sinio survey location contradicted the
northern boundary of M. tonkeana at Labuan Sore reported by Fooden [1969].
These field results and information from local people prompted us to examine
Raven’s original collection map (on loan from the Smithsonian to Dr. G.G. Musser
of the American Museum of Natural History). This map placed Labuan Sore at
latitude 0°37´S instead of 0°27´S, or in the approximate location of our survey
site of Avalua (Fig. 2).
The morphology of the Labuan Sore specimens was recently reevaluated by
J.W. Froehlich using a modified set of the diagnostic traits developed in this
study. Crown Hair Orientation was not included; character states for this fea-
ture were determined by the relationship between fur, skin, and the shape of the
skull in living monkeys, and were thus not comparable in museum specimens. In
addition, diagnostic features that were found to have a developmental compo-
nent (Cheek Whisker Contrast and Rump Patch Extent) in baseline populations
were not included. The Labuan Sore Mean Hybridity Index (LSMHI) was con-
202 / Bynum et al.

structed from the remaining three features: Gluteal Fields Extent, Ischial Callosity
Shape, and Shank Contrast [Bynum et al., in press]. The calculated LSMHI of the
three museum specimens indicated morphology between that of M. hecki and M.
tonkeana, and the animals collected in 1916 were significantly more M. tonkeana–
like than the current Avalua sample (Labuan Sore mean LSMHI 2.00, Avalua mean
LSMHI 1.33, n = 7; Mann-Whitney U-test statistic = 20.00, P = 0.03 with 1 d.f.).

DISCUSSION
Location and Description of the Hybrid Zone
We defined sets of morphological traits that were diagnostic for M. tonkeana
and M. hecki at the 95% or greater level and then located an area where animals
had intermediate or mosaic features. Hybridization as indicated by morphology
was detected between M. tonkeana and M. hecki (Fig. 2). The hybrid zone ap-
peared to be strongly centered at the Tawaeli–Toboli road (region 4). Only sev-
eral kilometers south of the TATO road, macaques sampled in Pangi-Binangga
(region 5) appeared to be almost entirely M. tonkeana–like. There were a few M.
tonkeana–like features in the Avalua population in a few kilometers to the north
(region 3), but 10 km further north in Sinio (region 2), we saw no indication of
anything other than M. hecki features in the population.
Morphology in the TATO area was not uniform. Animals in the western area
of the TAO region closely resembled the parental form of M. hecki, while mon-
keys in the far eastern areas were quite M. tonkeana–like. Variance in morpho-
logical scores was highest in the area of kilometers 50–59 along the TATO road,
but this result may have been influenced by the relatively large sample for this
area. Additional research must establish whether high morphological variation
adjacent to the road is also present in the areas adjacent to kilometers 40–50 of
the TATO road. In addition, we have used only morphological markers to define
the hybrid zone, while exhaustive research would also consider patterns of varia-
tion in genetic, environmental, and behavioral features.
Previously, it was suspected that the hybrid zone between M. tonkeana and
M. hecki might extend far up the isthmus that connects Central and North Su-
lawesi [Groves, 1980]. Based on the morphological data presented here, we esti-
mated the maximum dimensions of the hybrid zone as only 15 km from west to
east and 7.5 km from north to south. The apparent contradiction between our
data and the hybrid zone as described by Groves was resolved, however, when
fieldwork and examination of Raven’s collection map revealed that the actual
location of “Labua Sore” (Labuan Sore) was at latitude 0°37´S instead of 0°27´S
[Bynum et al., in press].
The only available evidence pertaining to the historical existence of the hy-
brid zone were the Labuan Sore museum specimens. We are hesitant to
overinterpret data from such a limited sample. However, if the Labuan Sore speci-
mens were representative of monkeys in that area in 1916, then the hybrid zone
has existed since at least 1916.

Effects of the TATO Road

The three Labuan Sore museum specimens exhibited more M. tonkeana traits
than the current Avalua sample, perhaps due in part to the fact that Labuan
Sore was approximately 1 km further south (i.e., toward M. tonkeana territory)
than our sampled Avalua locations. However, the possibility also existed that the
hybrid zone once extended further to the north than it does today.
 Sulawesi Macaque Hybridization / 203

The road from Tawaeli to Toboli was built from 1925–1930, 10 years after
Raven collected his specimens from Labuan Sore [Raven in Miller, 1915; Davis,
1976]. It will never be known for certain what relationship originally existed
between the hybrid zone and the placement of the TATO road or whether their
close cooccurrence is coincidence. The activities of humans may have created
secondary contact and hybrid zones between some taxa [reviewed in Hewitt, 1989],
including some primates [Fooden, 1964; Bernstein, 1966; Southwick & South-
wick, 1983]. In most cases, human activities involved habitat alteration such
that previously isolated taxa came into contact. In other cases, human activities
are thought to have modified or shifted existing zones [Hewitt, 1989].
The area between kilometers 40 and 50 on the TATO road was fairly heavily
disturbed at the time of this study. Agricultural land extended for up to a kilo-
meter an/or one ridge system on either side of the road. This created a mosaic of
habitat types hostile to macaques or from which they were actively chased away
by farmers. The deforestation in kilometers 40–50 may have acted as a semiper-
meable barrier to macaque movement from north to south and to a lesser extent
from east to west. Although it appears that habitat disturbance related to the
TATO road was limited until the road was paved in 1978, perhaps over decades
even this limited perturbation was enough to affect dispersal of macaques across
the area and pull the hybrid zone several kilometers south from Avalua/Labuan
Sore to lie basically coincident with the TATO road. If the hybrid zone once ex-
tended further to the north, disturbance associated with the road would have
disproportionately affected the ability of M. tonkeana to move from the south
into the hybrid area.
Barton and Hewitt [1985] and Hewitt [1988] have used the term tension
zone as a collective descriptor of clines in which hybrid unfitness is present as a
result of heterozygote disadvantage or maladapted hybrid recombinants. Tension
zones are maintained by a balance between dispersal and selection [Hewitt, 1988].
It has been observed that tension zones caused by genetic incompatibilities come
to rest in “density troughs” associated with habitat features such as valleys, riv-
ers, ridges, and roads [Barton & Hewitt, 1985; Hewitt, 1988]. However, it does
not follow that a hybrid zone spatially associated with an environmental feature
is necessarily a tension zone. These same environmental features can cause habitat
and density to vary for reasons unrelated to or in addition to the fitness of hy-
brids. The only way to determine if a hybrid zone is a tension zone or, in extreme
conditions, a hybrid sink is to measure selection against hybrids and dispersal of
parental and hybrid forms.

Origin of the Hybrid Zone

The origin of the hybrid zones by primary differentiation as opposed to sec-
ondary contact has long been a subject of controversy among evolutionary biolo-
gists. Endler [1977, 1982] demonstrated that a cline can be produced by vicariance
followed by differentiation and subsequent dispersal (secondary contact) or by
environmental heterogeneity producing spatially varying selection coefficients (pri-
mary differentiation) [Endler, 1982].
Most researchers working in hybrid zones have concurred that for a single
cline it is difficult to distinguish primary from secondary intergradation. How-
ever, concordant (or coincident) change in multiple characters across a hybrid
zone is “one of the stronger arguments in favor of origin by secondary contact”
[Hewitt & Barton, 1980]. Harrison [1990] viewed the distinction between zones
of primary differentiation and secondary contact as a probability function. As the
204 / Bynum et al.

number of coincident clines in nonlinked characters increases, so does the prob-

ability that the zone originated by secondary contact.
The data presented here do not permit the evaluation of strict concordance
of clines in the M. tonkeana/M. hecki hybrid zone. Traits evaluated in this study
were qualitative rather than quantitative, and cline width could not be mea-
sured with great precision. However, the clines for diagnostic morphological fea-
tures were broadly coincident within the M. tonkeana/M. hecki hybrid zone. This
should be considered preliminary evidence that the hybrid zone between the taxa
was a result of secondary contact. This is consistent with scenarios of allopatric
differentiation and subsequent recontact of Sulawesi macaques that have been
developed by several authors [Fooden, 1969; Hamada et al., 1987].

Taxonomic Status of M. tonkeana and M. hecki

Species definition and classification have long been contentious issues, com-
plicated by the sometimes competing aims of classifying biological diversity and
the identification of fundamental evolutionary units [Cracraft, 1989]. According
to the biological species concept (BSC), species are “groups of actually or poten-
tially interbreeding natural populations which are reproductively isolated from
other such groups” [Mayr, 1963]. Closely related to the BSC is the recognition
concept (RC), in which a species is a unit which shares a common fertilization or
specific-mate recognition system [Paterson, 1985]. Taken together, the BSC and
the RC can be viewed as reproductive species concepts, in that the role of gene
flow in either differentiating or maintaining cohesion within populations is em-
phasized [Endler, 1989; Templeton, 1989]. The BSC and RC have been criticized
for their inadequate treatment of asexual and parthenogenetic taxa. In addition,
the recognition of widespread and sometimes stable hybrid zones [Barton &
Hewitt, 1985; Harrison, 1990, 1993] between many plants and animals created
problems for both the BSC and the RC. Historically, species designation for hy-
bridizing taxa has often been a function of “whether the taxa were recognized as
distinct prior to the discovery of the hybrid zone” [Harrison, 1990].
Other species concepts emphasize factors in addition to reproductive conti-
nuity. According to the phylogenetic species concept (PSC) [Cracraft, 1983, 1989],
a species is an “irreducible (basal) cluster or organisms, diagnosably distinct from
other such clusters, and within which there is a parental pattern of ancestry and
descent.” The cohesion concept of species (CC) is a relatively new idea introduced
by Templeton [1989]. Of all species definitions, the CC explicitly acknowledges
the potential and varying roles of gene flow, genetic drift, and selection in pro-
ducing and maintaining species. Under the CC, a species is “the most inclusive
population of individuals having the potential for phenotypic cohesion through
intrinsic cohesion mechanisms” [Templeton, 1989.
The focus of this paper has been describing and interpreting the interaction
between M. tonkeana and M. hecki rather than establishing their taxonomic sta-
tus. Nevertheless, evidence indicated that M. tonkeana and M. hecki were two
distinct species according to the PSC, as the taxa were found to be diagnostically
different in a number of features [Cracraft, 1989]. (In the strict sense, it was
demonstrated that M. tonkeana and M. hecki were diagnosably distinct popula-
tions, but no attempt was made to establish the polarity of characters used. Thus,
while it was established that at a minimum M. tonkeana and M. hecki were
“evolutionarily significant units” [sensu Vogler and DeSalle, 1994], no phyloge-
netic hypothesis about their relationship was tested [see Bynum, 1995].) Hybrid-
ization between these two taxa may represent “the retention of a primitive
 Sulawesi Macaque Hybridization / 205

morphogenetic milieu” [Cracraft, 1989, p. 43]. Among macaques, where fertile

hybrids can be produced in captivity among many members of the genus [re-
viewed in Fa, 1989], the idea of “symplesiomorphic” hybridization between dif-
ferentiated forms may be particularly apt.
Proponents of the cohesion concept of species would also probably agree
that M. tonkeana and M. hecki should be considered two species. The taxa
were distinct over most of their geographic ranges. Hybridization appeared
restricted to a narrow area of concordant clines, although of course it was
possible that some alleles have introgressed far beyond the observed bound-
aries of the hybrid zone. There was evidence of historical depth for the hybrid
zone. Persistent differences in spite of hybridization argued that some compo-
nent of genetic or demographic exchangeability was not fully met by the hy-
bridizing populations and that separate species status was warranted according
to this framework [Templeton, 1989].
The relative narrowness of the hybrid zone, along with the probability that
contact between M. tonkeana and M. hecki was not new, suggested that there
was some sort of barrier to full admixture between the two taxa. Such a barrier
could be prezygotic, postzygotic, or environmental. It was necessary to take a
closer look at morphological, demographic, ecological, and behavioral features of
the hybrid population in order to address this issue. The results of this more
intensive study will be presented in forthcoming publications.
In conclusion, we note that because of its shape the isthmus has always
acted as a natural funnel between ranges of M. tonkeana and M. hecki. In recent
years, the funnel has been squeezed tighter due to forest habitat destruction
(Fig. 2). If further work is to be undertaken in the hybrid zone between M.
tonkeana and M. hecki, the time is now.

ACKNOWLEDGMENTS
This research was undertaken as one component of the Sulawesi Primate
Project, a collaborative and multidisciplinary effort to understand the evolution,
behavior, and ecology of Sulawesi primates. The work was sponsored by the In-
donesian institute of Sciences (LIPI) for allowing the work to be undertaken. In
Central Sulawesi, we acknowledge the assistance and support of the local office
of the Directorate of Forest Conservation and Nature Protection (PHPA), in par-
ticular Ir. Atang Setiawan, Mr. Abdoel Noerchayat, Mr. Rolex Lameanda, and
Mr. Yopiy Rumangkang.
This work would not have been possible without the guidance and support of
Dr. A.F. Richard, chair of E.L. Bynum’s doctoral committee, and the other mem-
bers of the committee: Dr. Carel van Schaik, Dr. Steven Beissinger, and Dr. Rob
DeSalle. Dr. David Higdon, Dr. Frances White, and especially Dr. Robert Dewar
provided invaluable assistance and guidance in data analysis and presentation.
We gratefully acknowledge the contribution of Dr. J. Froehlich in reexamining
the “Labua Sore” specimens at the Smithsonian Museum.
Financial support for this project was provided by a Fulbright-Hays Doctoral
Dissertation Research Abroad Fellowship, a National Science Foundation Doc-
toral Dissertation Improvement Grant (with A.F. Richard), the Wenner-Gren Foun-
dation for Anthropological Research, the Douroucouli Foundation, the Leakey
Foundation, the National Geographic Society, the Chicago Zoological Society, World
Wildlife Fund, the Williams Fund of the Department of Anthropology at Yale
University, Sigma Xi, the Enders Fellowship of Yale University, and American
Women in Science.
206 / Bynum et al.
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