Rank and grooming reciprocity among females in a mixed-sex group of captive hamadryas baboons

In a mixed-sex, captive group of hamadryas baboons (Papio hamadryas hamadryas) we investigated whether female grooming relationships are affected by their dominance ranks. Seyfarth's [1977] grooming for support model and Barrett et al.'s [1999] biological market model both predict that in primate groups where competition for monopolizable resources is high, grooming among females is based, at least partly, on the interchange of grooming for rank-related benefits, and that rank thus influences the distribution of grooming in females. Contrary to this prediction, our results show that despite the existence of a linear dominance hierarchy, rather strict dominance relationships, and high food-related aggression rates, grooming among female hamadryas baboons is not affected by rank and is only exchanged for itself. This is understandable since rank differences in our study group only result in differential access to limited, preferred food items that are not actively shared. Although some females are more likely to tolerate one another at the food pile, this tolerance is not determined by their grooming efforts and interchange of grooming for rank-related benefits does not occur. We conclude that female hamadryas baboons groom others in order to be groomed by them, which is supported by our observation that grooming reciprocity within a dyad increases when more grooming occurs in this dyad. Our results indicate that grooming is indeed a valuable commodity in itself, probably because of its stress- and tension-reducing effect. Based on our findings, the existing groom trade model is extended to include circumstances in which monopolizable resources are available but are not traded for grooming.     

Verreaux's eagles (Aquila verreauxi) as potential predators of hamadryas baboons (Papio hamadryas hamadryas) in Eritrea

In this paper we describe for the first time encounters of Verreaux's eagle (Aquila verreauxi) with hamadryas baboons (Papio hamadryas hamadryas) in the central highlands of Eritrea (15° 22′ N, 38° 58′ E, 2300 m). During 12 h of observation on 4 days, we observed four encounters of Verreaux's eagles with baboons, of which three can be classified as possible attacks. The baboons always responded with alarm calls. In three cases some immatures rushed to adult group members and clung to them, particularly to the adult male. Adult males threatened the eagle, and the whole group did not flee. The response of baboons towards smaller raptors like tawny eagles (Aquila rapax) and black kites (Milvus migrans) was very different. The adult baboons gave no alarm barks but scanned the raptors. The permanent presence of a pair of Verreaux's eagles in the home range of the baboons may represent a considerable predation pressure, at least for immature baboons. Am. J. Primatol. 47:61–66, 1999. © 1999 Wiley‐Liss, Inc.     

Conflict management by hamadryas baboons (Papio hamadryas hamadryas) during crowding: a tension-reduction strategy

Primates change a variety of behavioral responses during short-term exposure to crowding. Under crowded conditions, rates of aggression, submissive behavior, and affiliative behavior may increase or decrease. Different patterns of change among these three categories of response have been interpreted as various coping "strategies" for managing the increased risk of conflict under crowded conditions. Grooming is of particular interest because this behavior is known to have a calming influence on the recipient and could be used to manage conflict under tense situations. A captive group of nine hamadryas baboons (Papio hamadryas hamadryas) composed of two harems was observed under short-term crowding to determine whether this species adopts conflict-management strategies similar to those described in other primates. The aggression, submission, affiliation, and displacement activities of the six adults in the group (two males and four females) were recorded, and behaviors in their small indoor quarters was compared with baseline behaviors in the outdoor section of their enclosure, which had over 10 times more space. Repeated-measures analysis of variance (ANOVA) was used to compare behavioral rates during crowding with baseline rates recorded immediately after crowding and during matched controls collected at the same time of day as the crowding sessions. Aggression and submission rates did not change significantly across conditions. Huddling together and proximity increased during crowding, and females increased grooming of their harem male during crowding. Displacement activities (e.g., pacing and self-grooming) increased during crowding, but scratching, an indicator of anxiety in primates, did not. The pattern of behavior exhibited by this group conforms to an active "tension-reduction" strategy in which animals successfully reduce the higher risk of aggression during crowding.     

Use of palm trees as a sleeping site for hamadryas baboons (Papio hamadryas hamadryas) in Ethiopia

Hamadryas baboons sleep on cliffs throughout their range, and this can be attributed to the safety cliffs provide against predators in the absence of tall trees. In this paper, we report the first documented occurrence of hamadryas baboons sleeping in doum palm trees rather than on cliffs. Data derive from a study of hamadryas baboons at the Filoha site in lowland Ethiopia. During all-day follows, data were collected on travel patterns, band activity, and location. Variation in the baboons' home range was characterized using vegetation transects. We discovered that one band in this population, Band 3, occasionally slept in doum palm trees (Hyphaene thebaica). The palm tree sleeping site differed from other palm fragments in the baboons' home range in that it contained a higher density of palm trees. Possible factors influencing this unique use of palm trees as a sleeping site include access to palm fruit, avoiding contact with Afar nomads, avoiding sharing sleeping cliffs with other bands, protection from predators, and the lack of cliffs in a section of the baboons' home range. Evidence from this study suggests that the palm tree sleeping site is used because it affords better protection from predators than other palm fragments in an area of the band's home range that does not contain cliffs.     

Taste Preference Thresholds for Food-Associated Sugars in Baboons (Papio hamadryas anubis)

We aimed to determine taste preference thresholds for 5 food-associated sugars in Papio hamadryas anubis. In a 2-bottle preference test of brief duration (2 min) 4 subadult baboons significantly preferred concentrations as low as 10 mM sucrose, 20 mM fructose, lactose, and maltose, and 25 mM glucose over tap water. Presentation of suprathreshold sugar solutions led to marked concentration-dependent polydipsia. The results show that baboons are among the most sugar-sensitive nonhuman primates tested so far and, thus, support the assumption that Papio hamadryas anubis may use sweetness as a criterion for food selection.     

Distribution and Habitat Associations of Baboons (Papio hamadryas) in Central Eritrea

At least three diurnal primate taxa are still present in Eritrea, NE Africa: hamadryas baboons (Papio hamadryas hamadryas), olive baboons (Papio h. anubis) and grivet monkeys (Chlorocebus aethiops aethiops). However, information on status and distribution of primates and their habitats in Eritrea is outdated and incomplete. We conducted a primate survey, focussing on hamadryas baboons, to obtain data which will be integrated in a national wildlife management and conservation plan in Eritrea. We obtained information about the geographical distribution and abundance of baboons, their altitudinal range, habitat quality of their home-ranges, aggregation sizes at sleeping cliffs and predator presence. We described habitat quality via the Normalized Difference Vegetation Index (NDVI), a vegetation classification of Landsat MSS satellite data. Hamadryas and olive baboons are still present in Eritrea in ample numbers. Their geographical distributions in 1997 and 1998 did not deviate significantly from their historical distributions. An estimated 15,000 Papio hamadryas hamadryas lived in the 25,000-km² area of survey (0.58 baboons/km²). Population densities of hamadryas baboons in many parts of the survey area are higher than at Kummer's (1968) study site in Ethiopia. Hamadryas baboons live at all altitudes in four of five ecogeographical zones of Eritrea. Olive baboons replaced them in the western lowlands. Both baboon taxa tend to select better quality habitats, characterized by a higher normalized difference vegetation index (NDVI) than the average for the respective ecogeographical zones. Hamadryas baboons show a greater ecological plasticity than olive baboons, which are confined to riverbeds with extended gallery forest. By the end of 1999, a hybrid zone could not be confirmed.     

Group Composition and Adult Sex-ratio of Hamadryas Baboons (Papio hamadryas hamadryas) in Central Eritrea

During a survey of the geographical distribution and abundance of hamadryas baboons (Papio hamadryas hamadryas) in central Eritrea, we collected detailed demographic data on six bands at four sites in different ecogeographical zones. The proportions of age-sex classes within the six bands differed only with respect to juveniles. The general social organization of the Eritrean hamadryas baboons is similar to that reported for Ethiopia and Saudi Arabia. Eritrean hamadryas baboons live in a nested fission-fusion system, with one-male units as the basic social entity. Although the baboons were not provisioned, as they are in many places in Saudi Arabia, and habitat quality in Eritrea is lower than that in Ethiopia, sex-ratios and group composition corresponded more to those found in the Saudi Arabian population. Sex-ratios within the study population, in bands and also in one-male units were significantly more female-biased than in Ethiopian ones, and one-male units tended to be larger. Data from Eritrea suggest that these differences are due to a combination of a heavily fluctuating rainfall pattern and differential maturation of the sexes.     

The Socioecology of Network Scaling Ratios in the Multilevel Society of Hamadryas Baboons (Papio hamadryas hamadryas)

Multilevel or modular societies characterize a range of mammalian taxa, allowing social groups to fission and fuse in response to ecological factors. The modular society of hamadryas baboons has previously been shown to consist of 4 levels: troop, band, clan, and one-male unit (OMU). A recent study by Hill et al. (Biology Letters 4:748–751, 2008) revealed a mean scaling ratio across successive levels of multilevel societies of ca. 3; this was consistent across elephants, orca, geladas, and hamadryas baboons. Here we reanalyze the scaling ratio for hamadryas baboons with previously unavailable data from Filoha. Our analysis revealed a mean scaling ratio for hamadryas of 3.28 without data on the hamadryas clan layer of organization at Filoha, but a ratio of 6.17 with these data included. This discrepancy is due to the large clan and band sizes at Filoha yielding a larger than average gap between the OMU and the clan. Further analysis revealed subsets of OMUs within clans, suggesting a 5th level of society in this population. When this 5th layer of social structure is included in the analysis, the scaling ratio at Filoha is consistent with that of other hamadryas populations and other taxa. These results suggest that a consistent mammalian scaling ratio can be used to detect previously hidden levels of organization within societies and to predict their sizes in taxa for which detailed behavioral data are not available.     

Testing the function of reconciliation and third-party affiliation for aggressors in hamadryas baboons (Papio hamadryas hamadryas)

In social groups, agonistic conflicts can have different negative consequences. Several post-conflict interactions have been suggested as post-conflict management behaviors to mitigate those negative effects. In this study, we investigated the function of two post-conflict behaviors--reconciliation and aggressor-initiated third-party affiliation--on the aggressor's levels of post-conflict anxiety and aggression in a large colony of hamadryas baboons. We also examined variation in the aggressor's levels of post-conflict anxiety as a function of relationship quality between the opponents as predicted by the Integrated Hypothesis. We found that after conflicts hamadryas baboon aggressors showed increased rates of anxiety-related behaviors and that they were also more likely to be involved in renewed aggressive interactions. Although both reconciliation and aggressor-initiated third-party affiliation reduced the probability of receiving post-conflict aggression, only reconciliation reduced the rates of anxiety-related behaviors, suggesting that the aggressors' post-conflict anxiety might be owing mainly to the damage that the conflict causes to their relationship with the victim. Furthermore, aggressor's rates of post-conflict anxiety were higher after conflicts with individuals with whom they had a high-quality relationship, supporting the idea that levels of post-conflict anxiety mediate the occurrence of reconciliation depending on the quality of the relationship with former opponent as predicted by the Integrated Hypothesis.     

Early sexual maturity in male hamadryas baboons (Papio hamadryas hamadryas) and its reproductive implications

We present data on sexual maturity in young hamadryas baboon males (Papio hamadryas hamadryas) and its reproductive consequences in a large captive baboon colony. Hamadryas baboons live in a multilevel social system, with one-male units (OMUs) as the smallest social entity. Male leaders of OMUs are believed to monopolize matings within their OMUs; hence mating is believed to be polygynous and monandrous. In a captive colony of hamadryas baboons, we found evidence that young males less than 4 years old fathered at least 2.5% of 121 offspring born subsequent to vasectomy of all adult males, and males aged 4-5 years fathered at least 16.5% of the offspring. Additional evidence that these young males are able to sire offspring came from a morphological comparison of sperm from hamadryas males of different ages. The sperm of a 48-month-old hamadryas baboon were morphologically indistinguishable from viable sperm from adult males, whereas sperm from a 45-month-old male showed some aberrations. If successful copulations by adolescent males constitute a regular pattern even in free-ranging hamadryas baboons, a hamadryas male's chances to reproduce would not be limited to his role as an OMU leader as previously assumed, and a male's reproductive career would consist of two phases: the adolescent phase, and the OMU leader male phase.     

A baboon (Papio hamadryas) model of insulin-dependent diabetes

Over a period of four years, streptozocin has been used to induce diabetes in 10 baboons, all of whom are insulin dependent. We describe our experience with their husbandry, induction of diabetes, insulin therapy, metabolic control and growth rate. Streptozocin dosage of 60 mg/kg readily induces hyperglycemia with minimal hepatic or renal toxicity. Using a once daily injection of mixed short and intermediate acting insulins at a dosage of 2–4 U/kg, it is possible to maintain a degree of metabolic control similar to that attained in patients.     

Hormonal Correlates of Divergent Growth Trajectories in Wild Male Anubis (Papio anubis) and Hamadryas (P. hamadryas) Baboons in the Awash River Valley, Ethiopia

Comparative investigations of hormone concentration and pattern during ontogeny can offer insight regarding the evolution of growth trajectories. Anubis (Papio anubis) and hamadryas (P. hamadryas) baboons exemplify primate populations at a crucial stage of phylogenetic divergence. Though not reproductively isolated, the species are distinguished by consistent morphological, behavioral, and physiological differences, including trajectories of growth and maturation associated with divergent male reproductive strategies. As a step toward understanding the proximate causes of these differences, we tested several hypotheses regarding the relationship of growth-regulatory hormones and binding proteins (insulin-like growth factor-I, insulin-like growth factor binding protein-3 [IGFBP-3], growth hormone binding protein, and testosterone) to growth in several measurements. We collected samples (N = 559) across 13 field seasons, from 7 different social groups. Samples came from 398 different individuals. We sampled 285 once; 76, twice; 29, three times; 5, four times; and 3, five times. Although ages at peak hormone concentrations were not significantly different, concentrations of all hormones and binding proteins measured, except testosterone, were higher in hamadryas than in anubis. All factors measured correlated positively with growth in both species, and IGFBP-3 and testosterone in particular correlated significantly with growth in all measurements. Overall, our findings suggest a role for the growth hormone/insulin-like growth factor axis in producing distinctive patterns of growth in these species.     

Digit Ratios and Dominance in Female Baboons (Papio hamadryas and Papio ursinus)

A growing body of literature suggests that the ratio between the second and fourth digits of the hands (2D:4D ratio) is associated with exposure to prenatal sex hormones in a variety of animals including primates. Female baboons form dominance hierarchies composed of matrilines of related individuals, and the social mechanisms contributing to the structure of these hierarchies have been well studied. We here investigated the relationship between inferred prenatal androgen effects (PAE) and female rank in a captive troop of Hamadryas baboons (Papio hamadryas) with a typical social structure and three captive groups of chacma baboons (Papio ursinus) made up entirely of unrelated orphans. Low 2D:4D ratios (high inferred PAE) were associated with higher-ranking females and high 2D:4D ratios (low inferred PAE) with lower-ranking females in both focal species. This negative correlation between 2D:4D ratio and rank suggests prenatal androgens are linked with the maintenance of female ranks within matrilines in troops with a natural social structure and to the ranks acquired by orphan females.     

Ontogenetic Scaling of Fore- and Hind Limb Posture in Wild Chacma Baboons (Papio hamadryas ursinus)

Large-scale interspecific studies of mammals ranging between 0.04–280 kg have shown that larger animals walk with more extended limb joints. Within a taxon or clade, however, the relationship between body size and joint posture is less straightforward. Factors that may affect the lack of congruence between broad and narrow phylogenetic analyses of limb kinematics include limited sampling of (1) ranges of body size, and/or (2) numbers of individuals. Unfortunately, both issues are inherent in laboratory-based or zoo locomotion research. In this study, we examined the relationship between body mass and elbow and knee joint angles (our proxies of fore- and hind limb posture, respectively) in a cross-sectional ontogenetic sample of wild chacma baboons (Papio hamadryas ursinus) habituated in the De Hoop Nature Reserve, South Africa. Videos were obtained from 33 individuals of known age (12 to ≥108 months) and body mass (2–29.5 kg) during walking trials. Results show that older, heavier baboons walk with significantly more extended knee joints but not elbow joints. This pattern is consistent when examining only males, but not within the female sample. Heavier, older baboons also display significantly less variation in their hind limb posture compared to lighter, young animals. Thus, within this ontogenetic sample of a single primate species spanning an order of magnitude in body mass, hind limb posture exhibited a postural scaling phenomenon while the forelimbs did not. These findings may further help explain 1) why younger mammals (including baboons) tend to have relatively stronger bones than adults, and 2) why humeri appear relatively weaker than femora (in at least baboons). Finally, this study demonstrates how field-acquired kinematics can help answer fundamental biomechanical questions usually addressed only in animal gait laboratories.     

Testicular Size,Mating System,and Maturation Schedules in Wild Anubis and Hamadryas Baboons

We report body mass and testicular size in 258 anubis (Papio anubis or P. hamadryas anubis) and 59 hamadryas (P. hamadryas or P. h. hamadryas) baboons, live-trapped in Ethiopia. As predicted by theories of sexual selection by sperm competition, among hamadryas baboons, which are monandrous, fully adult males have absolutely and relatively smaller testes than those of comparable males among anubis baboons, which are polyandrous. Male hamadryas are also ca. 10% smaller in bodily mass as adults. The intertaxonal difference in adults is due entirely to the fact that in male anubis baboons, testicular and bodily mass continue to grow up to full adulthood–the age at which most males emigrate from their natal troop and initiate a confrontational breeding strategy among unrelated animals. By contrast, male hamadryas baboons, which are usually philopatric, attain adult body mass and testicular size as subadults. In both species, juveniles experience rapid testicular growth peaking in rate at ca. 12kg body mass, but testicular descent and growth starts earlier in hamadryas than in anubis baboons. Juvenile hamadryas baboons have relatively larger testes than their anubis equivalents, perhaps because male philopatry allows the mating strategy of male hamadryas baboons to be initiated during juvenile life and therefore permits some sperm competition between juveniles and adults.     

Gc subtypes identified by isoelectric focusing in Baboons (Papio hamadryas)

Summary Phenotyping for Gc variants by conventional electrophoresis in 835 Papio hamadryas baboons demonstrated a monomorphic population. Gc subtyping by polyacrylamide IEF gels, pH 4–6, on 394 of these baboons revealed the existence of two common alleles which we named Gc _Papio ¹ and Gc _Papio ² . Pedigree data confirmed the inheritance of a single locus, two allele system and the observed gene frequencies were 0.593 for Gc _Papio ¹ and 0.407 for Gc _Papio ² .