Distribution and possible intergradation betweenMacaca tonkeana andM. ochreata at the borderland of the species in Sulawesi

Morphological observations of pet and wild monkeys were made in the area that was inferred to be the borderland betweenMacaca tonkeana andM. ochreata in Sulawesi. Almost all individual monkeys could be classified into one of two species by their external characteristics. The possible borderland was estimated to extend from the La River in the east and to around Karaena River in the west. These two species may make contact in the forest in the western area of the borderland. Some external characteristics exhibited wide individual variations in the two species. Some monkeys originating from the borderland showed external characteristics that were intermediate between those of the two species. The possible intergradation between these two species is discussed in terms of the morphological variations found in the two species.     

Confirmation and location of the hybrid zone between wild populations of Macaca tonkeana and Macaca hecki in Central Sulawesi,Indonesia

Reports of hybridization between Macaca tonkeana and Macaca hecki were investigated in Central Sulawesi, Indonesia. We defined sets of morphological traits that were diagnostic for M. tonkeana and M. hecki and then located an areas where animals had intermediate or mosaic features. Hybridization as indicated by morphology was detected between M. tonkeana and M. hecki. The hybrid zone appeared to be strongly centered at the road that crosses the isthmus of Central Sulawesi from Tawaeli to Toboli. Macaques in this region were not morphologically uniform; 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 diagnostic morphological features were broadly coincident, suggesting that the hybrid zone originated by secondary contact. Analysis of three museum 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.     

The borderlands and possible hybrids between three species of macaques,M. nigra,M. nigrescens,andM. hecki,in the northern peninsula of Sulawesi

The borderlands between three species of macaques,Macaca nigra, M. nigrescens, andM. hecki, which live on the northern peninsula of Sulawesi were surveyed mainly via observations of pet monkeys kept by local people. The borderlands between these species could be delineated. Some monkeys of peculiar appearance and/or with mixtures of the external characteristics of two species were found in the borderlands between the respective pairs of species. However, such possibly hybrid monkeys were seen or originated only in the very limited areas where the species might come into contact with each other.     

Hematological values and parasite fauna in free-rangingMacaca hecki and theM. hecki/M. tonkeana hybrid group of Sulawesi Island, Indonesia

Hematological values and parasitological fauna of free-rangingMacaca hecki and the hybrid group betweenM. hecki/M. tonkeana of Sulawesi Island, Indonesia, were investigated. The hematological values, especially the red cell number (RBC), were lower than those of other macaque species, indicating that Sulawesi macaques are slightly anemic. Several parasites including Plasmodium sp., trombiculid mites, andTrichuris trichiura were identified. Although infection by Plasmodium was observed with considerable frequency, no clear relationship between its infection and the occurrence of anemia was found. Trombiculid mites and eggs ofAnatrichosoma sp. were detected in foci of the ears of most monkeys. The infection with a trombiculid mite is the first recorded occurrence in free-ranging wild Sulawesi macaques. Gastrointestinal parasites were identified from their eggs in fecal samples, where five species of nematoda and one trematoda species were found.     

Species differences of male loud calls and their perception in sulawesi macaques

Playback experiments were conducted to investigate interspecific discrimination of male loud calls in Sulawesi macaques. Loud calls of four macaque species living in Sulawesi (Macaca tonkeana, M. maurus, M. hecki, andM. nigrescens) and a control stimulus (an 8-sec frequency modulated sound) were played back to semi-free-ranging Tonkean macaques (M. tonkeana). A preliminary acoustic analysis indicated that the calls of these four species differ in some spectral and temporal features. In the playback experiments, Tonkean macaques responded in a similar manner to conspecific calls and calls of two other species,M. maurus andM. hecki. In contrast, animals responded more weakly to the call ofM. nigrescens and the control stimulus. Males responded more strongly than females to all stimuli, while females appeared to be more discriminating for species differences than males. Analyses on the acoustic features of loud calls suggested that high frequency, wide frequency range, and repetition of sound units at a high rate elicit quick responses from animals.     

Morphological studies on the Sulawesi macaques I: Phyletic analysis of body color

The body color of Sulawesi macaques was measured quantitatively and compared among the different monkeys. As a result, divergence models for extant Sulawesi macaques, withtonkeana as the starting point and fading as the sole direction of color change, were inferred as follows: (1) fading slightly on the upper half of the body—nigra, fading more on the proximal part of the body—nigrescens; (2) fading over the whole body—maura; (3) fading greatly on the legs—hecki; and (4) fading on the distal part of the body—ochreata, fading more over the whole body, including the proximal part of the body—brunnescens. The color changed progressively in the order of (1) through (4). The divergence model, excluding the position ofhecki (3), supports the speciation model ofFooden (1969). If the proto-Sulawesi macaques had a body color pattern similar to the livingnemestrina, darkening would have been necessary for the evolution of the Sulawesi macaques after their immigration, and it may have been acquired as an adaptation to the ground (forest floor) living nature of the Sulawesi macaques, together with influences deriving from the insularity and/or from the absence of predators.     

Origin and evolution of the sulawesi macaques 1. Electrophoretic analysis of hemoglobins

The monkeys on the island of Sulawesi (Celebes), Indonesia, comprise seven species ofMacaca, that isM. maura, M. tonkeana, M. hecki, M. nigrescens, M. nigra, M. ochreata, andM. brunnescens. Hemoglobins from 248 individuals of these seven species were analyzed by isoelectric focusing electrophoresis (IEFE) and by starch gel electrophoresis in the presence of urea (USGE). Eighteen phenotypes consisting of eight molecular types were identified by IEFE analysis. The speciestonkeana inhabiting the central part of the island revealed 11 phenotypes, while peripheral species such asnigrescens andbrunnescens carried only 3 and 2 phenotypes, respectively. On USGE, three α chains and three β chains were identified and named α1, α2, and α6, and β1, β3, and β5, respectively. The α1 chain has the same mobility as the α chains of other macaques, while the α2 chain is less positively charged than α1, and α6 is the least positive among these α chains. The α2 chain is widely distributed in the Sulawesi macaques as the major component. Four species,ochreata, tonkeana, maura, andnigrescens, carried the α1 and α6 chains as minor components. The electrophoretic mobility of β1 was the same as that of other macaques, while β3 and β5 were more positively charged and less positively charged than β1, respectively. All of the Sulawesi species had β3 in high or low gene frequencies and inmaura, tonkeana, andbrunnescens, this type was most abundant. β5 chain existed in the species of the northern peninsula, as the major type. The subordinate type was β3 innigra andnigrescens and β1 inhecki. On the other hand, β1 was most frequently observed inochreata.     

Macaca nigra on Bacan Island,Indonesia: Its morphology,distribution, and present habitat

We studied the monkeys inhabiting Bacan Island on the Maluku Archipelago, Indonesia. We compared them morphologically with Sulawesi black macaques (Macaca nigra).We also assessed their distribution on the island and on neighboring islands and the influence of human activity on them. We found that the Bacan black macaques are essentially similar to Sulawesi macaques; the variations in each locality are comparable. The monkeys probably inhabit the whole island or, at least, the lower forest thereon. Human activities — cultivation, logging, catching and killing monkeys — have not much affected their population. Thus they constitute a good stock of Macaca nigra,though the main population in Sulawesi is endangered.     

Reliability of the emotions profile index,primate form,withPapio hamadryas,Macaca fuscata,and twoSaimiri species

An evaluation of the reliability of the Emotions Profile Index (EPI), primate form, was carried out using two captive species of Old World monkeys (Papio hamadryas andMacaca fuscata) and two species (?) of New World monkeys (Saimiri sciureus andS. boliviensis). Observers, some familiar with the animals and some unfamiliar, rated members of the four groups at different times. Inter-rater reliability was high for most members of all species, but only when the observers were familiar with the animals. Assessments remained stable over at least one year.     

Origin and evolution of the sulawesi macaques 2. complete amino acid sequences of seven β chains of three molecular types

Seven β chains were identified as the typical molecular types carried by the seven species of Sulawesi macaques based on isoelectric focusing and urea starch gel electrophoresis. These β chains include the β3 chains ofmaura, tonkeana, nigra, andbrunnescens, β1 chains ofhecki andochreata and β5 chain ofnigra. The results of chromatography on cation-exchange and reversed phase columns and the amino acid compositions of the tryptic peptides suggested substitutions at the 9th and 13th amino acids from the N-terminal. Sequence analyses of these seven β chains from the N-terminal to the 18th amino acid and those of purified tryptic peptides from βT3 to βT15 by Edman degradation revealed the following facts: (1) the amino acid sequences of the β3 chains carried by the four species coincided with each other and as did those of the β1 chains of the two named species; and (2) the 9th and 13th amino acids were Lys and Thr in β3, Asn and Asn in β1, and Asp and Thr in the β5 chain, respectively. These three β chains are related with each other by at least two-base changes. The evolution of the β chains of the Sulawesi macaques was inferred to be as follows. (1) The β3 chain might have been dominant β chain in the past among Sulawesi macaques, since peripheral species separately carried this chain; (2) the β1 and β5 chains might have derived from a “missing link” because of more than two-base substitutions between β3 and β1 and between β3 and β5; (3) eight other macaque species, including the lion-tailed macaque (M. silenus), bear Asn and Thr at these two positions, while the Barbary macaque (M. sylvanus) has Thr and Thr; and (4) thus, if the parsimonious rule is followed, the type with Asn-Thr is the most plausible “missing link,” since only the Asn-Thr type can combine these five β chains by minimum one-base change. Two genetic events are postulated in the evolutionary process of the Sulawesi β chains: the first Lys-Thr type (β3) was distributed over the whole island, and next Asn-Thr, the common type in other macaques, produced Asn-Asn (β1) and Asp-Thr (β5).     

Japanese macaques form a cross-modal representation of their own species in their first year of life

We tested whether infant Japanese macaques (Macaca fuscata) have a cross-modal representation of their own species. We presented monkeys with a photograph of either a monkey or a human face on an LCD monitor after playing back a vocalization of one of those two species. The subjects looked at the monitor longer when a human face was presented after the monkey vocalization than when the same face was presented after human vocalization. This suggests that monkeys recall and expect a monkey’s face upon hearing a monkey’s voice.     

Handedness found in a wild group of moor monkeys Macaca maurus in the Karaenta Nature Reserve, South Sulawesi,Indonesia

Hand preference of wild moor monkeys Macaca maurus was investigated in food reaching situations at the Karaenta Nature Reserve, South Sulawesi, Indonesia. The frequency picking up sweet-corn grains to take into the mouth by either hand of monkeys was counted directly at the feeding ground where sweet-corn grains were scattered for monkeys. Among the 20 monkeys examined, 8 were fight-handed, 8 were left-handed, and 4 were ambilateral. The results indicated the prevailed hand preference on individual level but not either trends of left- or right-hand preference on population level. The trend toward a higher proportion of left-handed monkeys found in Japanese and rhesus monkeys was not found. Some other characteristics found in moor monkeys are discussed in comparison with those previous findings in Japanese and Tibetan macaques in order to evaluate variations within the genus Macaca.     

Rapid identification of white-Engelmann spruce species by RAPD markers

Fragments of random amplified polymorphic DNA (RAPDs) were used as markers to distinguish Picea glauca (Moench) Voss (white spruce) and Picea engelmannii Parry (Engelmann spruce). These species and their putative hybrids are difficult to differentiate morphologically and are collectively known as interior spruce. Four oligodeoxynucleotide decamer primers showed species-specific amplification products between white spruce and Engelmann spruce. These fragments are highly conserved among seed lots and individual trees of each species from diverse geographic origins. The consistency and reproducibility of these species-specific amplification products were tested in more than two amplification reactions. Therefore, RAPD markers can provide genetic markers for easy and rapid identification of the specific genetic entry of these spruce species and their reported putative hybrids. According to the frequencies of the species-specific RAPD markers, it is possible to estimate the hybrid fraction, indicative of true introgression between the two species. These results are useful for quick identification of both species and their hybrid swarms at any stage in the sporophyte phase of the life cycle, for determining the occurrence and the magnitude of introgressive hybridization in an overlap zone between the two species, and for certification purposes in operational re-forestation and tree-improvement programs.     

Vertical distribution of wild Yakushima macaques (Macaca fuscata yakui) in the western area of Yakushima Island, Japan: Preliminary report

A census of wild Yakushima macaques (Macaca fuscata yakui) was carried out in a 23-km² area of the western coast of Yakushima Island, Japan. We analyzed the census data to investigate changes in monkey distribution associated with the vertical distribution of vegetation. In the lowland coastal zone of 0–300 m above sea level (a.s.l.), 4.8 troops and 62.4–99.8 monkeys are estimated to have existed per km². In the mountainside zones of 300–900 m a.s.l., the troop density decreased to 1.3–1.6 troops/km². Since there was no difference in size between the coastal and mountainside troops, population density should decrease with altitude to about 30–36 monkeys per km². On the other hand, 2.4 troops and about 36 monkeys were estimated to have inhabited per km² in the mountain summit zone of 900–1,323 m a.s.l. Nature Conservation College     

Reconciliation in three groups of lion-tailed macaques

We studied postconflict behavior in three captive groups of lion-tailed macaques (Macaca silenus). After a conflict, we monitored the aggressee as the focal individual during a 10-min postconflict period and made control observations the following day on the same individual. Selective attraction between former opponents occurred in the first minutes of the postconflict period. The conciliatory tendency was relatively high, about 40%. Although no specific behavior was used to reconcile, postconflict contacts were especially intense and a rich repertoire of affiliative patterns was exihibited. With regard to the rate and form of reconciliation, lion-tailed macaques resemble Sulawesi macaques, which belong to the same phyletic lineage. We also discuss the possible interrelations between conciliatory patterns and other characteristics of social organization.     

Discrimination of macaques by macaques: the case of Sulawesi species

A series of work by the first author have demonstrated that many macaque species show a visual preference for the pictures of their own species when the monkeys actively press a lever to see the pictures. We expanded this study to Sulawesi macaques kept as a pet by local people with slight modification. All seven species of Sulawesi macaques were passively exposed to a variety of colored slides of Sulawesi macaques. The experimenter recorded the duration of visual fixation onto the pictures. Male monkeys of all the seven species clearly watched the pictures of their own species for longer duration than those of the other species. Such visual preference suggested that the seven Sulawesi macaques discriminate each other species and, thus, they may not be integrated into fewer number of species. This visual preference may work to prevent overall intergradation of the Sulawesi macaques who sometimes have hybrid zones only in limited areas. This preference was in general weaker in female monkeys. In one species,Macaca ochreata, females actively avoided to see the pictures of conspecifics. These results may be related to how female monkeys interact with other individuals.     

Behavior, Diet, and Movements of the Sulawesi Crested Black Macaque (Macaca nigra)

We present the first field study of activity budgets, diet and ranging patterns of the Sulawesi crested black macaque, Macaca nigra, one of seven macaque species endemic to Sulawesi, Indonesia. We studied three crested macaque groups, ranging in size from 50 to 97 individuals, for 18 months in the Tangkoko-DuaSudara Nature Reserve, North Sulawesi. They spent 59% of the day moving and procuring food, especially fruits, and 41% of the day resting and socializing. Their diet is composed of more than 145 species of fruit (66% of observed feeding bouts), vegetative material (2.5%), invertebrates (31.5%), and occasional vertebrate prey. Group differences were more pronounced than seasonal or diurnal differences. Specifically, the largest group moved farther during the day, moved at a faster and more uniform rate, ate less fruit, rested more, and socialized less than the smaller groups did. The largest group had the largest home range, but it included less primary forest and more disturbed habitat than the ranges of smaller groups. There are individual differences in activity budgets of adult males and females in time spent moving, resting, feeding, and socializing that may reflect differences in reproductive strategies of males versus females. The behavior of large juveniles is more similar to that of adults than to that of small juveniles. Daily movements and use of home range are correlated with diet. Macaques moved shorter distances as the proportion of time spent feeding on fruit increased, and the top four dietary items accounted for most of the variance in entry into hectare blocks of home range.