External characteristics and associated developmental changes in two species of Sulawesi macaques,Macaca tonkeana andM. hecki,with special reference to hybrids and the borderland between the species

The degree of intergradation between two species of Sulawesi macaques,Macaca tonkeana andM. hecki, was studied by examining the diagnostic external characteristics of more than 100 monkeys kept as pets by natives. Two possible hybrid monkeys were found and both originated from the borderland between the two species, located in the most proximal region of the northern peninsula of Sulawesi. The previously postulated wide area of integradation between the two species at the possible contact zone was, however, not recognized, and typical examples oftonkeana orhecki were found to be present on the two sides of a narrow “hybrid” zone which was defined by direct observations. Furthermore, despite considerable individual variations, we were able to allocate most monkeys to one or other of the species. Each of ten external characteristics of the members of both species more or less encompassed the individual variations, but may undergo changes with the development of the monkeys. The mechanisms of reproduction of hybrid monkeys and the maintenance of differences between the species are discussed.     

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.     

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.     

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.     

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.     

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).     

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.     

Reappraisal of the taxonomic status of the cranium Stw 53 from the Plio/Pleistocene of Sterkfontein,in South Africa

A fossil skull, Stw 53, from the Plio/Pleistocene of Sterkfontein, in South Africa, has been referred toHomo habilis Leakey, Napier, andTobias, 1964. Reappraisal of its putative hominine affinity reveals a closer resemblance toAustralopithecus africanus Dart, 1925. The skull, as reconstructed, is too small forH. habilis; with no indication of brain expansion overA. africanus; has a facial angle outside the hominine range, but identical with that ofA. africanus; and whose teeth are not elongated but display buccolingual expansion. Although it was found in the same strata (Member 5) as stone tools, there is no causal connection. It has been dated faunistically at 2–1.5 my BP, but due to an unconformity it is suggested that it could be older. In spite of its late date, Stw 53 shows no intermediate characters which could support a trend towardsH. habilis orA. robustus Broom, 1938. It may, therefore, represent a relict population ofA. africanus.     

Intergroup encounters in wild moor macaques (<Emphasis Type="Italic">Macaca maurus</Emphasis>)

We studied intergroup encounters among moor macaques at the Karaenta Nature Reserve, South Sulawesi, Indonesia. Group B has been observed on the basis of individual identification since 1988. We analyzed 85 encounters between members of Group B and members of neighboring groups from September 1990 to November 1998. The average frequency of intergroup encounters was 0.035/hour. Neither the presence of females in estrus nor rainfall had an effect on encounter frequency. Behaviors of moor macaques during intergroup encounters differed from those of Japanese macaques. In moor macaques, no intergroup interactions with body contact were observed during encounters, and females never directed aggression toward members of different groups. The present study did not confirm the prediction of the model ofvan Schaik (1989). Extension of the existing models is required to explain the difference in female dominance styles among macaques by socioecological factors.     

Latitudinal,taxonomic, sexual,and insular determinants of size variation in pigtail macaques,Macaca nemestrina

Regression analysis demonstrates that body and skull size are correlated with latitude in Sundaic (Macaca nemestrina nemestrina)and Indochinese (M. n. leonina)pigtail macaques. These two contiguously distributed subspecies are unusual in that they exhibit opposite latitudinal gradients of increasing size— M. n. nemestrinabecomes larger in a southerly direction,while M. n. leoninabecomes larger in a northerly direction in accordance with the usual pattern of ecogeographic variation seen in other sympatric macaques and mammals (Bergmann ’s rule). The difference in clinal size gradients is one more of a series of characters which delineate these two macaques as natural biological populations. However, since the size gradients converge on a narrow zone along the Thai-Malay Peninsula, the use of size as a character to evaluate genetic and morphological intergradation is equivocal when population variation is considered. The third subspecies, the Mentawi Island pigtail (M. n. pagensis),is an endemic,insular isolate differentiated from the Sundaic pigtail from similar latitudes in terms of its small size. Thus, latitude, insularity, and taxonomic differentiation all affect size variation in the pigtail macaques;also,although the data are not definitive, there is the suggestion that the degree of sexual dimorphism may be an additional covariate of latitude and/or body size.     

Even more new taxa from South and East Anatolia II

10 new Turkish taxa are described:Arenaria eliasiana, A. sivasica, A. monscragus, A. angustifolioides; Campanula lycica; Scutellaria orientalis subsp.tortumensis; Stachys choruhensis, S. tundjeliensis; Calamintha caroli-henricana; Aristolochia rechingeriana, the latter two species named in honour ofKarl Heinz Rechinger;Allium vuralii. Dedicated to Prof. DrKarl Heinz Rechinger on the occasion of his 80th birthday. For part I see Pl. Syst. Evol.154, 111–128.     

Positional behavior and social organization of the philippine tarsier (Tarsius syrichta)

This study provides preliminary data on the positional behavior and habitat use of the Philippine tarsier,Tarsius syrichta, which was studied in its natural habitat on the island of Leyte. The locomotor behavior ofT. syrichta is quite comparable to the other species of tarsiers but frequency of use of different postures and support types does seem to distinguish this species from the others.Tarsius bancanus appears to be the most specialized, using vertical clinging postures the most frequently and sitting the least;Tarsius dianae uses vertical clinging the least frequently and sitting the most, andT. syrichta is intermediate. The Philippine tarsier has a small home range similar to that ofT. spectrum. On the other hand, this species likely does not have the small family groups typical of Sulawesi tarsiers but instead uses a noyau system likeT. bancanus. This data supports some apsects ofNiemitz’s (1977) model of anatomical and behavioral specialization among tarsier species.     

Isolation of a new flavonol glycoside and its effects on the blue color of seed coats ofOphiopogon jaburan

From the blue seed coats ofOphiopogon jaburan, a new flavonol glycoside was isolated as needles and determined to be kaempferol 3-O-β-d-galactoside-4′-O-β-d-glucoside (OK-2) by UV and NMR spectral analyses. OK-2 and kaempfrol 3, 4′-di-O-β-d-glucoside (OK-1), which was detected previously, in the blue seed coat were present in a molar ratio of about 13:7. OK-2 was newly found as a factor causing the blueing effects on ophionin which is a main anthocyanin in the blue seed coats. The mixture of 4.8×10⁻³ M OK-2 and 2.5×10⁻³ M ophionin in Mcllvaine's buffer solution (pH 5.6) showed stable blue color, and the absorption spectrum of the mixture showed two absorption peaks and a shoulder in visible reasion, coinciding with that of the fresh blue seed coat. The effect of ophionin and OK-2 co-pigmentation on the blue color of seed coat ofO. jaburan was discussed.     

Rapid shifting of foraging pattern by Yakushima macaques ( Macaca fuscata yakui ) in response to heavy fruiting of Myrica rubra

We describe short-term changes in foraging behavior by wild Yakushima macaques (Macaca fuscata yakui),which inhabit a warm-temperate broad—leaved forest on Yakushima Island (30°N, 131°E), Japan. Rapid changes of dietary composition, activity budget, and range use by the monkeys occurred from May to June, apparently associated with changes in the availability of the fruit of Myrica rubraBefore the fruit ripened, monkeys spent less time moving and more time feeding on many species of leaves, which accounted for 40% of feeding time. However, when M. rubrabegan to ripen, they fed intensively on the fruit, which accounted for three-fourths of feeding time,though the activity budget remained unaffected As fiuit of M. rubradecreased,the monkeys fed more on the fruit of other species and on insects, and spent more time moving at higher speeds. There marked shifts in foraging pattern occurred within only two months. In terms of moving cost and dietary quality,Yakushima macaques shifted their foraging pattern according to the availability of M. rubrafrom a “low-cost, low-yield” strategy to a “low-cost, high-yield” strategy, and then to a more costly strategy. The ability to make such rapid shifts in foraging pattern may allow the macaques to effectively use the highly variable food supply within their small range.     

Insectivory and its significance to langur diets

Earlier studies on Hanuman langurs (Presbytis entellus) around Jodhpur found them entirely vegetarian (Mohnot, 1974;Winkler, 1988). However, recent observations in the open scrub forests of Jodhpur reveal that scale insects may constitute a regular part of their diet and that insectivory is particularly prevalent in the monsoon months July – September. The insect eating in this habitat seems to support the “Energy/nutrient maximization” hypothesis ofHamilton andBusse (1978).     

Protein variation,taxonomy and differentiation in five species of marmosets (genusCallithrix Erxleben, 1777)

The electrophoretic patterns of 15 protein systems codified for 20 genetic loci were investigated using horizontal electrophoresis. A total of 150 blood samples, from five species of the genusCallithrix were analyzed. Polymorphic variation was observed in 10 out 20 loci analyzed. The genotypic distributions are in Hardy-Weinberg equilibrium. The average heterozygosity (H) varied from 1% to 5%, similar to those observed for other Neotropical primates. The genetic distance coefficients revealed a phylogenetic separation of these species into two groups: (1) “argentata” (C. humeralifer andC. emiliae); (2) “jacchus” (C. jacchus, C. penicillata, andC. geoffroyi). This arrangement is according to the taxonomic arrangement proposed byHershkovitz (1977),de Vivo (1988), andMittermeier et al. (1988). The results in each group are compatible with the subspecies values recorded for the Platyrrhini. These values showed that:C. humeralifer andC. emiliae are subspecies ofC. argentata;C. jacchus, C. penicillata, andC. geoffroyi are subspecies ofC. jacchus. These results also suggest thatC. j. geoffroyi is the “jacchus” group taxon, most similar genetically to the “argentata” group.     

Risultati della lotta biologica contro ilDacus oleae Gmel. a mezzo dell'Opius concolor Sz. siculus Mon.

Summary The Author reports on the essays of artificial biological control of the olive fly (Dacus oleae Gmel.) made by himself andGenduso with the specific parasiteOpius c. siculus Mon. found in Sicily. The Authors followingDelanoue's technic have bred theseOpius in laboratory on larvae ofCeratitis capitata Wied; so they were able to release: in 1961 — 1000Opius in a small olive plantation near Palermo; in 1962 — 24000Opius in the island of Pantelleria (Sicily); in 1963 — 40000Opius in the island of Salina (Eolic Islands-Sicily); in 1964 — 180000Opius also in the island of Salina. The releasedOpius are ever reproduced, reducing theDacus pest. From the agricultural standpoint the results have been satisfactory and the Author hope to release a great number ofOpius in the olive plantation of all Eolic islands, in 1965.

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Nota:Fischer (1958, 1963) ha pubblicato un lavoro sugliOpius della Regione etiopica, nel quale, a parziale modifica del precedente guidizio di sinonimia, dichiara perl'Opius siculus Mon.: ?sarebbe meglio considerarlo una razza del concolor? (?am besten als Rasse von O. concolorSzépl. aufzufassen?).     

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.