Mating patterns,mate choice,and birth season heterosexual relationships in free-ranging rhesus macaques

Birth season adult heterosexual nonkin relationships of 50 free-ranging female rhesus macaques (Macaca mulatta) in two social groups at Cayo Santiago, Puerto Rico were examined using focal follow (289 hr) and ad lib data. Eighty-eight percent of subjects had at least one relationship characterized by particularly high frequencies of spatial proximity, grooming, or both. These were designated “friendships.” Males intervened in aggressive interactions more frequently on behalf of Friends than non-Friends. Female aggressive support of males was extremely rare. Higher-ranking males experienced more friendships than lower-ranking males. High-ranking females had higher-ranking Friends than low-ranking females. Older females had higher-ranking Friends than younger females. Females groomed high-ranking Friends more than they were groomed by them, whereas they groomed low-ranking Friends less than they were groomed by them. In one social group, high-ranking females were more likely than low-ranking females to groom their Friends more than they were groomed by them. Males were more responsible than females for spatial proximity maintenance in 9 of 14 Friend dyads for which sufficient data were available. Neither male nor female dominance rank affected responsibility for proximity maintenance in Friend dyads. Eight of 24 females had friendships with males with whom they had completed copulations during their conception peri-ovulatory period of the preceding mating season. Two of 19 females completed peri-ovulatory copulations with Friends during the following mating season. Friendship was not correlated with either of two demonstrated female mate choice indicators: (1) proximity maintenance during estrus; or (2) cooperation with male “hip-grasp” courtship attempts. Males directed “muzzle-up” courtship signals at lower rates toward Friends than toward non-Friends. These and other investigators' results indicate that (1) protection from aggression is the primary benefit to female rhesus macaques of birth season heterosexual relationships; (2) the most effective protectors are in greatest demand as Friends; and (3) friendship has no effect or an inhibitory effect on mate choice in this species. Benefits to males of friendships were not apparent from this study but may include coalitional support against lower-ranking males.     

Attributes and validity of dominance hierarchy in the female pigtail macaque

A group of six unrelated female pigtail macaques,Macaca nemestrina, of the same age, was studied in captivity over a period of five years. The animals were observed under six different situations, following changes in the social composition of the group. The following social interactions were recorded: withdrawal, attack, threat, presentation, mount, and grooming. Although behavioural rates varied for each period, their distribution network was remarkably consistent. Each subject could be assigned a rank, which remained stable over the six periods. From an analysis of the number of dyads in which one of the two animal significantly performed both paired behaviours more than the other animal (external validity), behaviours clustered into two main groups: (1) attacks, threats, and withdrawals; and (2) mounts, presentations, and grooming. Quantitative methods were performed to standardize the degree in which linearity, stability, reciprocity, and idiosyncrasy of the interactions under study may account for social variability. When considering these properties, the behaviours clustered in the same two groups (“agonistic” and “affinitive”). A method to describe the dominance style ofMacaca nemestrina was proposed, which can easily be replicated for comparisons with females of other primate species and in different conditions.     

Social behavior of the emperor tamarin in captivity: Components of agonistic display and the agonistic network

Agonistic behavior was studied longitudinally for 16 months in an intact family groups of captive emperor tamarins (Saguinus imperator subgrisescens) using methods from quantitative ethology and social network analysis. A motivational analysis of the components of agonistic display revealed the relative strength of each component along a continuum from strongly dominant to strongly subordinate. Tabulations of exchanges of strongly dominant and strongly subordinate components in interactions among the tamarins revealed an agonistic network (“dominance hierarchy”) that approached, but did not quite reach, the ideal state of a transitive order (“linear dominance hierarchy”). The frequency with which individual tamarins long called and scent marked was not closely correlated with their position (“dominance rank”) in the agonistic network. Instead, individuals undergoing change in status long called and scent marked frequently, irrespective of their rank.     

Rank and age related feeding strategy observed through field experiments in the Koshima group of Japanese macaques

The feeding process of Japanese monkeys on soy beans which were scattered over a sandy beach on Koshima islet was studied. Younger monkeys were able to pick up more beans when 8 kg of beans were divided and given two times (“two times feeding”) than when the whole amount (8 kg) of beans was given at one time (“one time feeding”). The effect of saturation of the food intake capacity in younger monkeys at the first feeding in “two times feeding” did not appear at the second feeding one hour later. The minutely intake of soy beans (feeding speed) for each age class was analyzed. The decline of feeding speed in adult females after the peak in “one time feeding” was not related to the decline in density of beans on the ground, and this decline was caused by saturation of their food intake capacity. Adult females were divided into four classes according to their dominance rank order: high, lower-high, higher-low, and low classes. The total amount of intake in “one time feeding” was far larger in the high class than in any of the other classes. The total amount of feeding in the first feeding of “two times feeding” increased in accordance with rise in the dominance rank class, and there was no relation to rank and total feeding amount in the second feeding of “two times feeding.” Differences existed in the process of feeding between the rank classes. The feeding speed of the low class was as high as that of the high class on the curve of minutely intake, while the low class stopped feeding much earlier than the high class. The lower-high class displayed a low feeding speed, and stopped feeding the latest. The order of the duration to stay and to feed in the feeding area was lower-high > high > higher-low > low, and this order did not change under the three different feeding conditions, “one time feeding,” and the first and second feedings of “two times feeding.” Adolescent females tended to stay the longest duration in the feeding area among all age classes. Both the lower-high class females and adolescent females had an unstable social status in the Koshima group, and their social status affected their feeding behaviors. The feeding behaviors were similar in attitude depending on social status, and are considered to be maintained for a fairly long time. The feeding strategy of the lower-high class, in staying a longer duration in the artificial feeding area, and departing later, may be effective under the artificial feeding conditions, but it may be a bad strategy in a natural habitat where the food is not so clumped as in artificial feeding, and where choice of other food patches is possible. The above results agree well with previous reports for the Koshima group, indicating that the rank of the lower-high class females was unstable (Mori et al., 1989), and that their reproductive success was low (Watanabe et al., 1992).     

Reproductive patterns in mantled howler monkeys: Estrus,mate choice and copulation

The present study was undertaken to evaluate non-random mating patterns in two groups of mantled howler monkeys in two tropical dry forest habitats. Sexual dimorphism, female estrus stage, male dominance rank, sexual solicitations and copulations were assessed. Males are significantly larger than females, but female weight varies more than male weight. The length of female estrus cycles is comparable in both habitats, but females in the more strongly seasonal habitat demonstrate greater estrus synchrony relative to their numbers. Males solicit potential mates more frequently than females, a pattern explained by the relatively high rate of sexual solicitation by high-ranking males. Females in “peak” estrus solicit “alpha” males, while females in other stages of estrus solicit males equally by rank. Intersexual aggression occurs rarely, and “forced copulations” are attempted but, apparently, are unsuccessful. Sexual solicitations by “alpha” males and “peak” estrus females are most likely to lead to copulation, and “alpha” males are more likely to copulate than “gamma” males. In general, latencies from first solicitation to copulation are expensive in time, especially for high-ranking males. Estimated annual reproduction success favors high-ranking males, and results indicate that male and female mating behavior is mutually coordinated and controlled.     

The evolution of “egalitarian” and “despotic” social systems among macaques

Recent studies of captive macaques have revealed considerable inter-species differences in dominance styles among females. In “egalitarian” species such as stumptail (Macaca arctoides) or tonkean macaques (M. tonkeana), social interactions are more symmetrical and less kin-biased than in “despotic” species such as Japanese (M. fuscata) or rhesus macaques (M. mulatta). Field observations of moor macaques (M. maurus), close relatives of tonkean macaques, suggest that tolerance during feeding characterizes their egalitarian dominance style in the natural habitat. Although it has been proposed that communal defense against other groups may be the main selective force in the evolution of egalitarian dominance style among females, few field data support this prediction. A game theory analysis showed that both an “egalitarian” strategy and a “despotic” strategy are possible evolutionarily stable strategies (ESS) under certain conditions. The difference in dominance styles might reflect the difference in ESS. This means that an egalitarian dominance style can emerge without strong between-group contest competition. A phylogenetic comparison among macaques suggests that despotic dominance styles very likely evolved from egalitarian dominance styles. In the future, primate socioecological studies should pay more attention to the evolutionary history of each species.     

The effects of dominance rank and group size on female lifetime reproductive success in wild long-tailed macaques,Macaca fascicularis

Demographic changes were recorded throughout a 12-year period for three social groups ofMacaca fascicularis in a natural population at Ketambe (Sumatra, Indonesia). We examined the prediction that females' lifetime reproductive success depended on dominance rank and group size. Average birth rate was 0.53 (184 infants born during 349 female years). For mature females (aged 8–20 yr) birth rate reflected physical condition, being higher in years with high food availability and lower in the year following the production of a surviving infant. High-ranking females were significantly more likely than low-ranking ones to give birth again when they did have a surviving offspring born the year before (0.50 vs 0.26), especially in years with relatively low food availability (0.37 vs 0.10). Controlled comparisons of groups at different sizes indicate a decline in birth rate with rroup size only once a group has exceeded a certain size. The dominance effect on birth rate tended to be strongest in large groups. Survival of infants was rank-dependent, but the survival of juveniles was not. There was a trend for offspring survival to be lower in large groups than in mid-sized or small groups. However, rank and group size interacted, in that rank effects on offspring survival were strongest in large groups. High-ranking females were less likely to die themselves during their top-reproductive years, and thus on average had longer reproductive careers. We estimated female lifetime reproductive success based on calculated age-specific birth rates and survival rates. The effects of rank and group size (contest and scramble) on birth rate, offspring survival, age of first reproduction for daughters, and length of reproductive career, while not each consistently statistically significant, added up to substantial effects on estimated lifetime reproductive success. The group size effects explain why large groups tend to split permanently. Since females are philopatric in this species, and daughters achieve dominance rank positions similar to their mother, a close correlation is suggested between the lifetime reproductive success of mothers and daughters. For sons, too, maternal dominance affected their reproductive success: high-born males were more likely to become top-dominant (in another group). These data support the idea that natural selection has favored the evolution of a nepotistic rank system in this species, even if the annual benefits of dominance are small.     

A study on the social structure and dispersal patterns of hamadryas baboons living in a commensal group at Taif,Saudi Arabia

Three levels of hamadryas social structure—the one male unit (OMU), the band, and the troop—have been observed at all sites studied, but a fourth—the clan—has been observed at only one site, Erer-Gota, Ethiopia, during a longitudinal check of the dispersion of identified individuals. The clan is important since it appears to provide the basis for male philopatry, although comparative data is needed from other sites to confirm this. We studied a huge commensal group of hamadryas baboons (over 600 animals) in Saudi Arabia. We put ear tags on baboons between 1998 and 2004 and analyzed social structure, relying on the interactions of these tagged animals by focusing especially on their dispersal patterns from OMUs. OMU membership tended to be looser than that of the Ethiopian hamadryas. Females tended to shift between OMUs on an individual basis in our study group, whereas the collapse of an OMU was a major occasion of adult female transfer in Ethiopia. We found neither stable bands (a “band” in our study group was defined as a regional assemblage of OMUs) nor clans that lasted for several years. Some OMUs moved and transferred into neighboring areas over both the short and long term. Further, some post-adolescent males appeared to move out of the study area. The ratio of adult females in an OMU in our study group was larger than for any other documented study site, and this may be the reason for enhanced female transfer between OMUs. A large proportion of the adolescent females showed no clear membership to OMUs, and no “initial units” (commonly observed in Ethiopia) were discernible. The ease with which young males acquired adult females at the study site must have disrupted the formation of a clan, a “male-bonded society.”     

Social relations between adult males and females of Japanese monkeys in the arashiyama B troop

Some dyads of Japanese monkey adult males and females show remarkable spatial proximity and frequent exchanges of social behaviors. It is suggested that some kind of “affinity” exists between them. Females obtain much unilateral benefit from “proximity effects”; even lowranking females can dominate high-ranking females as long as they stay nearby their “affinitive” males. Males acquire female followers in return. Mating relations and female mother-daughter relations play important roles in forming new “affinitive relations.” Once monkeys have formed “affinitive relations,” however, they seldom mate with each other, as if they were kin-related. Therefore, the acquisition of female followers appears inconsistent with a male's strategy for reproducing many genes in the next generation. This study was financed partly by the Cooperative Research Fund of the Primate Research Institute, Kyoto University. The outline of this paper has already been published inTakahata (1980b).     

Taxonomic reassessment ofBolonia Meunier, 1886 (trace fossil) based on new material from the type area in Boulonnais, northern France

The ichnospeciesBolonia lata Meunier, 1886 is a straight, bilobate trace fossil with a somewhat heart-shaped outline in cross-section. It has previously been reported from various localities mainly from the Jurassic by different authors under various names such asGyrochorte, Scolicia, orProtovirgularia. Because none of these previous determinations fully agrees with the trace fossils’ morphology, the ‘forgotten name’ is reintroduced, and due to a lack of type material a neotype from the type area of Boulonnais, France is proposed. The ichnogenus is currently represented only by one ichnospecies,B. lata Meunier, 1886. Diagnostic criteria ofBolonia and morphologically related trace fossils such asScolicia, Gyrochorte, Bichordites, “Taphrhelminthopsis nelsoni”, andProtovirgularia are discussed, the synonymy ofB. lata is provided and finally a discussion on the possible producer ofBolonia is given.      

Male Mating Tactics in Captive Rhesus Macaques (Macaca mulatta): The Influence of Dominance, Markets, and Relationship Quality

Male mating success in a multimale–multifemale group can depend on several variables: body condition, dominance, coalitions, “friendship,” or an exchange of services for mating access. Exchange patterns may also be determined by market effects or social relationships. We studied the mating tactics of males in a captive, multimale–multifemale group of rhesus macaques and the resulting patterns of mating and paternity to determine the influence of dominance rank, mating markets, and relationship quality on their mating tactics. Male rank was positively related to the total number of copulations and the number of mating partners, but did not explain male mating distribution completely. Moreover, male fertilization success was not related to male rank. Males did not exchange grooming for mating access on the same day and neither the supply nor the rank (as a proxy for quality) of receptive females affected the amount of male grooming, suggesting that market effects did not explain male mating access. However, there was a positive correlation between long-term grooming patterns of both males and females and mating access, indicating that social relationships were important for male mating access. Paternity data revealed that these social relationships were also important for male reproductive success. We conclude that both male rank and male–female “friendship” determined male mating access in these rhesus macaques, but that “friendship” was more important in determining paternity, emphasizing the importance of intersex social bonds in male mating success in multimale primate societies.     

Mating Resets Male Cricket Aggression

An animal’s motivational state can significantly impact its behavior. We examined the effects of mating on the aggression of male Acheta domesticus crickets. Pairs of males were allowed to establish dominance and subordinance and were then physically separated. Subordinate males were then allowed to either copulate with a female or to have chemo-tactile contact with, but to not copulate with, a female. Less than 15 min after separation, all male pairs engaged in a second agonistic encounter. Subordinate males that copulated with females were significantly more aggressive toward their dominant partners than un-mated subordinate males. Many mated subordinates became dominant. Allowing a subordinate male to contact, but not copulate with, a female had a similar effect, suggesting that chemo-tactile cues from the female are sufficient to elicit this change in aggression.     

Behavioural and physiological correlates of personality in greylag geese (Anser anser)

Personality means suites of correlated behavioural traits, also referred to as “behavioural syndromes” or “personality dimensions”. Across animal taxa similar combinations of traits seem to prevail, which may have proximate foundation in common neuroendocrine mechanisms. Hitherto, these have been rarely studied in intact social settings. We investigated personalities of greylag goose males from a free-roaming flock that shows complex social relationships. In connection with our longitudinal study on the consistency of behavioural and physiological responses to multiple challenges, we asked whether and how single, personality-related behavioural traits correlate with each other to form personality dimension(s). We tested whether these dimensions were related to physiological characteristics that previously showed limited plasticity (heart rate (HR), baseline and stress-induced excreted immuno-reactive corticosterone (BM), and testosterone metabolites levels) and, furthermore, to age, body measures, and dominance rank. Principal-components analysis based on behavioural variables revealed two factors: 51.1% of variability was explained by “aggressiveness” and a further 19.1% by “sociability”. “Aggressiveness” comprised correlated measures of aggression, subordinance, boldness, vigilance, and proximity to the mate. This “aggressiveness” positively correlated with stress-induced BM levels, the HR increase during aggressive interactions, and with dominance rank, which may suggest proximate and functional contingencies of this personality dimension.     

Influence of ovule perforation,plant growth regulators,andl-glutamine on in vitro growth of immature peach embryos

Summary Ovule perforation technique and media components (plant growth regulators andl-glutamine) were tested on in vitro growth of immature (<3 mm) embryos of “Springcrest” and “Earligrande” peaches. Ovule perforation was 2 to 4 times more effective in promoting embryo growth than leaving ovules intact.l-Glutamine (400 mg·liter⁻¹) promoted an increase in growth but could not be used with indole-acetic acid plus kinetin because an antagonistic effect on embryo growth occurred. The use of these exogenous plant growth regulators did not increase embryo growth over in vivo growth.     

Restudy and reassignment ofDentalium antiquum Goldfuss, 1841 (Middle Devonian)

Dentalium antiquum is one of two moderately well-known Devonian “scaphopods” in the German literature. Examination of the type material and a few specimens in other institutions indicates more individual variability than is to be expected in species of the molluscan class Scaphopoda. The species is transferred with question toColeolus Hall, a fossil presumed to be a calcareous “worm” tube.      

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.     

The functions of status in the mantled howler monkey,Alouatta palliata Gray: Intraspecific competition for group membership in a folivorous neotropical primate

In the dominance hierarchies of adult male and female mantled howler monkeys (Alouatta palliata Gray), high-ranking individuals are young adults; intermediate-ranking individuals, middle-aged adults; and low-ranking individuals, old adults. This relationship reverses the trend observed in most group-living animals and is previously unreported for this species. A limiting supply of palatable leaves may create intense intraspecific competition for group membership which, it is hypothesized, has resulted in this rare pattern of hierarchical relations. It is shown that individual-level selection is sufficient to explain the evolution of this apparently “altruistic” status system, though other mechanisms are assessed. The energetic constraints imposed by a folivorous diet appear to restrict the expression of aggression to “ritualized” forms. Two groups of monkeys in two different habitats were studied and the rates of appeasement and aggressive behavior were found to be higher in the more “stressful” forest. The latter group, however, displays significantly more behavioral patterns entailing low or intermediate energy expenditure.     

Decline in energy storage in the Antarctic minke whale (<Emphasis Type="Italic">Balaenoptera bonaerensis</Emphasis>) in the Southern Ocean

The annual trend in energy storage in the Antarctic minke whale was examined using catch data from all 18 survey years in the Japanese Whale Research Program (JARPA). Regression analyses clearly showed that blubber thickness, girth and fat weight have been decreasing for nearly 2 decades. The decrease per year is estimated at approximately 0.02 cm for mid-lateral blubber thickness and 17 kg for fat weight, corresponding to 9% for both measurements over the 18-year period. Furthermore, “date”, “extent of diatom adhesion”, “sex”, “body length”, “fetus length”, “latitude”, “age” and “longitude” were all identified as partially independent predictors of blubber thickness. The direct interpretation of this substantial decline in energy storage in terms of food availability is difficult, since no long-term krill abundance series is available. However, an increase in the abundance of krill feeders other than minke whales and a resulting decrease in the krill population must be considered as a likely explanation.     

Dominance and feeding competition in captive rhesus monkeys

The feeding behavior of 16 adult female rhesus monkeys living in three captive social groups was observed. Estimates of relative food intake, feeding rate, and location of feeding in relation to food sources were compared between females of different dominance ranks. Higher-ranking females had greater access to feeding sites and were supplanted or threatened less frequently while feeding than subordinates. However, no consistent differences in estimates of total intake were found between females of high and females of low rank. The effects of dominance on feeding behavior were most pronounced in the group receiving the least food relative to estimates of overall group nutritional requirements. Higher-ranking females, both over the long term and during the study period, tended to produce more surviving offspring. The effects of dominance on reproductive performance appeared to be less related to food intake than to competitive and aggressive interactions, potentially resulting in higher levels of stress for subordinates.     

Benefits and Costs of Dominance in the Angelfish Centropyge bicolor

Social groups are often structured by dominance hierarchies in which subordinates consistently defer to dominants. High‐ranking individuals benefit by gaining inequitable access to resources, and often achieve higher reproductive success; but may also suffer costs associated with maintaining dominance. We used a large‐scale field study to investigate the benefits and costs of dominance in the angelfish Centropyge bicolor, a sequential hermaphrodite. Each haremic group contains a single linear body size‐based hierarchy with the male being most dominant, followed by several females in descending size order. Compared to their subordinate females, dominant males clearly benefited from disproportionately high spawning frequencies, but bore costs in lower foraging rates and greater aggressive defence of their large territories. Within the female hierarchy, more dominant individuals benefited from higher spawning frequencies and larger home ranges, but displayed neither higher foraging rates nor spawn order priority. However, dominance in females was also linked to aggressiveness, particularly towards immediate subordinates, suggesting that females were using energetically costly aggression to maintain their high rank. We further showed by experimentally removing dominant females that the linear hierarchy was also a social queue, with subordinates growing to inherit higher rank with its attendant benefits and costs when dominants disappeared. We suggest that in C. bicolor, the primary benefit of high rank is increased reproductive success in terms of current spawning frequency and the prospect of inheriting the male position in the near future, which may be traded off against the cost of aggressively defending rank and territory.