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MANDRILLS & SAVANNA BABOONS.
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Examines social organization of two simian primate baboon species. Demographics, sex-specific size, appearance, life expectancy, habitats.... More...
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Paper Abstract:
Examines social organization of two simian primate baboon species. Demographics, sex-specific size, appearance, life expectancy, habitats.

Paper Introduction:
The purpose of this research is to examine the social organization of two simian primate baboon species, the mandrill (Mandrillus sphinx) and savanna baboons (Papio cynocephalus anubis). The plan of the research will be to set forth the context in which social structures of these species assume significance and then to discuss demographical features of each species, notably sex-specific size, appearance, sexual configuration and behavior, life expectancy, and both natural and artificial habitats. The interplay of environment and behavior and the implications of nonhuman primate physical health and behavior for human health and behavior suggest in general terms why the study of primates has significance in the modern period. In particular, evidence of primate social structure is suggestive because it demon

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Individual and social behavior in a captive troop of mandrills(Mandrillus sphinx). According to Tuttle (1987), aging baboons "become moreinclined to groom each other rather than to play, and well before they areadults, they often enter into another, more permanent, kind of male-malerelationship" (p. Smuts, R. Maledominance and genetically determined reproductive success in the mandrill(Mandrillus sphinx). Dixson, A. Parasitology,112, 489-497. According to Stammbach, most grooming occurs among members of thesame OMU or at least same "clan." Social status within the group and sexappear to be the most important factors of grooming behavior, although sizeof troop may also be significant. J., Bossi, T., & Dixson, A. Wickings, E. (1984). L. (1992, December). Thenatural habitat of baboons is the equatorial African rain forest. This is consistent withStammbach's discussion of "interband relations" observed among mandrills,which may share multiple-group feeding areas (1987, p. Primates, 28,289-3 8. A reasonable inference appears tobe that complexity implies a standard of social organization to which thebaboon population as a whole tends to adhere. The principal type of food gatheringengaged in by mandrills and savanna baboons alike is foraging (Westergaard,1992; Lahm, 1986), although baboons are generally recognized as omnivorous. Animal Behavior, 5 , 1667-1682. Males thatachieve the greatest tumescence in a group appear also to achieve thegreatest success in mating with and also guarding their mates fromincursion by other males; the ability to guard appears positively relatedto group dominance (Dixson, Bossi, & Wickings, 1993). Primates. D., Whiten, A., & Strum, S. 66), which areof the genus Papio. Lahm, S. Rogers, M. (1995). The first explanation, connected with the sternum rub, is that SMreinforces male-dominance (sexual) hierarchies in the troop. Mandrill in Gabon's rain forest--ecology,distribution and status. F. Napier, J. Primates, 22, 2 6-22 . (1996).Intestinal parasite burden in five troops of olive baboons (Papiocynocephalus anubis) in Gombe Stream National Park, Tanzania. To see whythis is the case, it is useful to cite Stammbach's observation (1987, p.112) that mandrills are among those species that form one-male units(groups), known as OMUs. Kudo, H. Aggressive male behavior, however, does not imply that aggression =success. C. P. Typically, both mandrills and savanna baboons are found in the wild intwo west African coastal equatorial countries, Cameroon (north of theequator) and Gabon (bisected by the equator), with natural features thatrepresent a "mosaic" of rain forest and savanna topography (Rogers &Others, 1996). (1988). S. In B. Blom, A., Alers, M. Folia Primatologica, 61, 115-122. (1993, October). M. Diet and habitat preference of Mandrillus sphinx inGabon: Implications of foraging strategy. A., Fontaine, B. 1 3 ) says that an assertion of "synonymy" for Mandrillus and Papio "hasnot found much support" in the field. (1991). 297-3 ). Mellen, J. E. Further, dominant femaleshave increased access to mating and feeding, and their offspring have agood chance of survival. Among mandrills, grooming does not appear to be well studied in thewild, owing to difficulties of access cited earlier. E., Abernethy, K. Primates of the world.New York: Blandford Publishing. (1988). Consider Stammbach's note--made prior to extensive observation andreportage in the literature of observations of mandrills in the wild--that"estrous females are mated nearly exclusively by their OMU leaders" (1987,p. Dominance rank, resourceavailability, and reproductive maturation in female savanna baboons.Behavioral Ecology and Sociobiology, 33, 313-318. Strum, S. In particular, evidence of primate social structure issuggestive because it demonstrates strategic response to practicalenvironmental conditions and points to systematic response to andinteraction with individual psychosocial units of which a given primatecommunity is composed. T., Feistner, A. J., White,L. Wickings, E. However, social behavior ofOld World Monkeys is not easily decoded, and experts are likely to continueto disagree on its meanings. Hoshino and others found that, in addition toOMUs, mandrills in the wild comprise multimale groups, with both solitarymales and subgroups of OMUs part of the mixture. The flamboyant appearance of the male mandrill appears to haveimplications for the sociobiology of the species. Inparticular, this behavior demonstrates that vocalized communication is adominance-hierarchy strategy that can be set beside sex-specific behavioras a factor of social-group analysis. Kudo alsoviews the patterns of vocal communication between groups of mandrills as anindex of their relative group openness, compared to the dynamics observedamong Theropithecus gelada baboons (Kudo, 1987). (1986). The plan of the research willbe to set forth the context in which social structures of these speciesassume significance and then to discuss demographical features of eachspecies, notably sex-specific size, appearance, sexual configuration andbehavior, life expectancy, and both natural and artificial habitats. Bercovitch, F. Field work on baboons shows the complexity of relationships in socialunits, dependent as they are on baboon thought processes, which are noteasily understood by human observation. The size of a mandrill troop may vary. 112-2 ). Acknowledging that reproductive behavior is an area ofcontroversy among scholars who disagree on whether male competition forfemale mates is partly a feature of secondary sex characteristics andsexually predatory behavior, Dixson, Bossi, and Wickings (1993) cite thedifference between so-called fatted and nonfatted males, also called large-rumped and slim-rumped, a reference to genital tumescence (Wickings &Dixson, 1992). According to Harrison (1988), mandrills may cluster in populationsthat appear to have limited scope to their habitat, compared to other wider-ranging savanna-dwelling simian species in the equatorial region, includingPapio cynocephalus anubis. Annual birth patterns ofsavanna baboons (Papio cynocephalus anubis) over a ten-year period atGilgil, Kenya. Social relationships andritualized greetings in adult male baboons (Papio cynocephalus anubis).International Journal of Primatology, 11, 147-172. Preston-Mafham, R., & Preston-Mafham, K. As alpha malesage and decline in rank or are otherwise displaced by more aggressivefatted males, they become marginalized and isolated as nonfatted/slim-rumped males from a social point of view. F., Bossi, T., & Wickings, E. Wickings and Dixson (1992) report observing accelerated puberty,testosterone, and testes development in dominant male mandrills. In particular,patterns of social dominance hierarchy have been observed in multimalemandrill groups, both in terms of more or less ordinary social interactionand in terms of sexuality dominance and control of females. Electrophoresis, 16, 1678-1683. J. (Ed.). (1993). M. B., & Strum, S. One reason for taxonomic controversyappears to be that mandrills are especially elusive in the wild and do noteasily present themselves for study (Harrison, 1988; Rogers, Abernethy,Fontaine, Wickings, White, & Tutin, 1996). (1992, November). Dixson, Bossi, andWickings (1993, pp. (1987). 16 ). The interplay of environment and behavior and the implications ofnonhuman primate physical health and behavior for human health and behaviorsuggest in general terms why the study of primates has significance in themodern period. Savanna baboons have beenobserved as far east as Kenya and Tanzania as well, although underrelatively controlled conditions (Packer, Collins, Sindimwo, & Goodall,1995; Bercovitch & Harding, 1993). Physiology & Behavior, 52, 9 9-916. A second(and logical) feature of observation was the sighting of both group/socialand solitary males. Mammalia, 52, 57-74. The mandrill, whichcan grow to the size of a giant dog and which is considered a quadruped(Napier, 1974), is also traditionally considered a savage or ferociousbaboon with highly developed sensory capabilities that affect groupbehavior (Phillips, 1975) and with an apparently "fearsome dental weaponry"(Tuttle, 1987, p. 117). Stammbach, E. There is evidence of dimorphism betweenadult males and females as well as between different categories of adultmales. The natural habitats of mandrills and savanna baboons differ. The mandrill appears to be among the largest physical type of OldWorld Monkey, weighing up to 3 kilograms but ranging between 21 and 18 kg(Stammbach, 1987; Napier, 1974). According to Kudo (1987), grunting behavior of mandrills is anindex of strong socialization patterns within multimale troops. Reproductive cycles also appear to be partly a function of availablefood supplies and other environmental factors, particularly among savannababoons. Tuttle, R. Scent marking in mandrills, Mandrillus sphinx.Folia Primatologica, 57, 42-47. As explained by Bercovitchand Harding, the diversity of food supplies influences fertility patterns.Another view is offered by Packer, et al., who interpret variable femalereproductive behavior in savanna baboons at a National Park in Tanzania insociological terms. J. Indeed, as reported by Wickings, Bossi & Dixson (1993),the DNA analyses of one mandrill troop observed in an animal preserve inGabon over five years indicated that two dominant males (one ascending indominance and one descending in dominance) were responsible for all infantbirths in the group. (1993, December).Reproductive success in the mandrill, Mandrillus-sphinx--correlations ofmale-dominance and mating success with paternity, as determined by DNA-fingerprinting. B., Watanabe, J. Westergaard, G. A., Sindimwo, A., & Goodall, J. (1987). First, they acknowledge that, like males, femalesmanifest dominance hierarchies in their group. Journal of ComparativePsychology, 1 6, 398-4 3. J. Smuts, B. One eight-month-long research investigation of captive mandrills,comprising one adult male, two females, and three young, cites a great manysimilarities in general behavior between mandrills and previously studiedbaboons (Papio) while also noting mandrill-specific facial expressions andbody language (Mellen, Littlewood, Barrow, & Stevens, 1981). Citing DNA data from the same basic mandrill group studied byWickings, Bossi, and Dixson, Wickings (1995) also reports evidence of 75percent avoidance of incestuous breeding, either because of the presence ofmore than one dominant male or because of the timing of maturation offemales as well as patterns of sexual avoidance on the part of certainselected females. There are two possibleexplanations for SM, which takes the form of sniffing a tree, then rubbingthe chin on it. This is consistent with Strum's finding (1987, p. The reproductive and therefore mating cycle of baboons isannual/seasonal (Dixson, Bossi, & Wickings, 1993). 14, pp. Apes of the world. 1 days in the life of a mandrill hordein the Lope Reserve, Gabon. 525-6) cite Darwin's reference to Mandrillus sphinx asthe brightest all mammals, with the species' sexual selection being basedon vibrance of hair color, plus blue and reddish coloration of (hairless)genitalia and face. According to Bercovitch andStrum (1993), the onset of puberty and willingness to mate can be explainedin terms of group sociology in general and group stress in particularbrought on by limited access to food and water. J. J., & Dixson, A. Not all behavior among baboons is aggressive and hierarchical. (1985). Comparative investigation of different species ofbaboons in all these areas, meanwhile, may be useful in refining anunderstanding of habitat and survival requirements of the species and maypoint in the direction of actions and policies that might have the effectof making highest and best human use of natural resources, revealingoptimal strategies for possible human intervention in primate populations,whether in captivity or the wild, and aiding in the construction ofinvestigations of human psychological, social, and physical development,including but not limited to an understanding of evolutionary origins ofhunting, socialization, and other behavior in early hominids. A third feature was slightly more complex. Fay, J. D., Littlewood, A. (1987, July). Preliminary reporton the grouping of mandrills (Mandrillus sphinx) in Cameroon. Primates, 26, 161-167. D. F. That same study revealed fairlycomplex social configurations among the various troops. In this regard, Bercovitch and Strum (1993) and Bercovitch andHarding (1993) separately report the same basic 1 -year study of a colonyof savanna baboons in Kenya, finding a positive relationship between femalereproductive behavior, environmental factors, and social organization.While conception as such appears to be a function of rainfall, food, andwater availability, the savanna baboons are otherwise not confined toseasonal reproductive patterns of behavior (i.e., copulation withoutconception). Fatted (tumescent) males are also said to be "fullycolored," while nonfatted males are considered "pale" (Rogers & Others,1996, pp. Packer, C., Collins, D. 114). Chicago: Encyclopaedia Britannica. Feistner, A. (199 ). This is consistent with the earlier observationthat alpha males in the same mandrill troop may be rising and descending.Thus, the persistence of troop cohesiveness and survival could beattributable to its ability to back away from confrontation at appropriatetimes, with both opportunistic and guarded mating coexisting in a unit atany given time. S. Other sources include Mandrills and drills in thedesignation of baboon, as species of all Old World/African monkeys that arenot anthropoid apes (Phillips, 1975). Genetic self-management in a captive colony ofmandrills (Mandrillus sphinx) as revealed by DNA minisatellitefingerprints. Priority of accessand grooming patterns of females in a large and a small group of olivebaboons. Meanwhile, there is a good deal of hunting of mandrills inparticular in the area, also a consequence of increased human penetrationand development of the habitat areas. Sambrook, T. C. Fatted males essentially function in the manner of "Type-A"personalities--more group-dominant, more flamboyant in appearance, and moresuccessful with females than the latter, in both sexual activity andpaternity success. Wrangham, & T. Both mandrills and savanna baboons (some of which are called olivebaboons on account of their coloring) are considered species of baboonsmore generally described as Old World Monkeys and included in the familyCercopithecidae (Preston-Mafham & Preston-Mafham, 1992, p. The purpose of this research is to examine the social organization oftwo simian primate baboon species, the mandrill (Mandrillus sphinx) andsavanna baboons (Papio cynocephalus anubis). Certain behavior that is not overtly sexual may nevertheless be anindex of social structure in mandrill populations and appears to beconnected to the fairly highly developed sensory capabilities of themandrill. New Britannica 3 (Vol. T. (1992). T.Struhsaker (Eds.). The reason for this may be the irregulardistribution of food sources, notably fruits, insects, and selected smallanimals, for mandrills in rain forest areas (Lahm, 1986; Kudo & Mitani,1985). Baboon. Testicular function,secondary sexual development, and social status in male mandrills(Mandrillus sphinx). Primates, 34, 525-532. For example, according to Sambrook,Whiten, & Strum (1995), savanna baboons who groom are likely to be lower onthe social hierarchy than those who are groomed; most grooming is same-sex. C. ORYX, 22, 218-228. Rain-forest-dwelling mandrills inhabit the ground,not the trees, although they are more likely to climb trees to reach foodthan savanna baboons are (Lahm, 1986). (1975). M. Harrison, M. Primates, 25,295-3 7. In recent years, sparse populations of Papio cynocephalusanubis have been counted as far north as the Central African Republic,which borders Cameroon on the east (Fay, 1988). S. There is some controversy on how to classify mandrills and baboons.Some sources differentiate Mandrillus sphinx, which manifests distinctivesphincteric (taut) facial marking around the nostrils, altogether from"baboons proper" (Preston-Mafham & Preston-Mafham, 1992, p. 61ff; Phillips,1975). New record of predatory behavior by themandrill in Cameroon. Undoubtedly the sensory features of mandrills areremarkable among lower primates. New York: McGraw-Hill. American Journal of Primatology,11, 9-26. Funk & Wagnalls New WorldEncyclopedia. According to Stammbach,birth may occur between December and April (1987, p. A good deal of evidence has been cited of connection betweenpsychosocial and psychobiological behavior in the mandrill malepopulations, with social rank a definite factor in group and individualencounter whether or not coloration fosters sexual success. O. W., &Barnes, K. This combination of factorscreates special challenges for any conservation and preservation measureson one hand (Blom & others, 1992) and the role that mandrills play indistributing seeds throughout their habitat range on the other (Lahm,1986). (1995).Reproductive constraints on aggressive competition in female baboons.Nature, 373, 6 -63. Wickings refers to this pattern of reproductive behavioras genetic self-management, over a period of nine years in which anoriginal colony of six mandrills increased to 84. Forest monkey populations in the Central AfricanRepublic: The northern limits. Primates in Gabon--current status and distribution.ORYX, 26, 223-234. There is also a human factor. However, Kudo and Mitani(1985) report a relative absence of dominance behavior between mandrills ina feeding situation that had involved cooperative killing of an antelope,and a similar pattern of predatory feeding in savanna baboons. T., & Tutin, C. The principal observations made byKudo demonstrate that calls between members of a troop can be classifiedaccording to age and sex and between subgroups. 1 14-1 31). On the other hand, female mating competition hasto be set beside the observation of negative effects; Packer and others(1995) observe increased likelihood of miscarriage during pregnancy fordominant females. C., & Stevens, V. Napier (1974)describes the cycle in terms of female reproductive periods: copulation,gestation, birth, and lactation, with females rejecting copulation untillactation and weaning of infants are completed. Chicago: University ofChicago Press. (1987). Vocal calls between females of different subgroups or males andfemales of different subgroups demonstrate the presence of socialinteraction in general, while certain vocal grunts between males of thesame group demonstrate dominance-hierarchy interaction in that group inparticular and the presence of one leader male. Adult male mandrills are roughly twice as large as adult females, andmale physical coloration is much more ornate than female physicalcoloration. J.(1981). Mandrillsare distinctive among baboon subspecies in that they appear to prefer todwell in the rain forest (Kudo & Mitani, 1985); savanna baboons, as thename implies, are more at home on grassland plains, subject to theavailability of water. The study of vocal communication of wildmandrills in Cameroon in relation to their social structure. B., & Harding, R. Hoshino, J., Mori, A., Kudo, H., & Kawai, M. G. The challenge remains how to analyze the lack of seasonality with theobserved irregularity of reproductive activity. New York: Funk & Wagnalls. (1992). Seyfarth, R. Journal of Zoology, 231, 563-574. Wickings, E. J. 114). Desert, forest and montane baboons: multilevelsocieties. F. One four-year longitudinalobservation of mandrills in Gabon identified troops numbering from 15 to 95(Hoshino, Mori, Kudo, & Kawai 1984). The secondexplanation is that SM reinforces habitat consciousness, keeping members ofthe troop from ranging too far away from home and getting lost. Object manipulation and the use oftools by infant baboons (Papio cynocephalus anubis). R. One view, however, is that the ferocitycharacterization, particularly attributed to adult male mandrills, ismisleading. A., & Woolhouse, M. C. T. This dimorphism, manifest as coloration, body build, and secondarysex characteristics, appears to have implications for the social structureand function of mandrill groups; colonies of baboons as a class arecommonly referred to as troops (Mellen & Others, 1981; Mueller Graf,Collins, & Woolhouse, 1996; Smuts & Watanabe, 199 ). (1996). For purposes of this research, andbased on the evidence of literature that identifies and analyzes themseparately, mandrills (Mandrillus) and savanna baboons (Papio) will beconsidered as separate subspecies of Old World Monkeys (FamilyCercopithecidae). Ranging behavior for mandrills has beenidentified as far north as southeastern Nigeria (bordering Cameroon on thenortheast) and as far south and west as the Zaire (Congo) River (Preston-Mafham & Preston-Mafham, 1992). As Blom, Alers, Feistner, Barnes, &Barnes, 1992) point out, the nearly two dozen primate species identified inGabon may in future years be squeezed out on account of lumber-industryactivities, which involve the creation of logging roads to remote areas andas a side effect the destruction or fragmentation of living space, henceredistribution of primate species throughout a given geographical area andpotential for decline of species population. Amongyounger males that do not appear to be dominant or among adult males whosedominance is giving place, mating success is infrequent and opportunistic,engaged in with unguarded females who then remain unguarded in the group bythe male(s) mating them (Dixson, Bossi, & Wickings, 1993). (1993). Scent-related behavior is described by Feistner (1991), who seessocial instincts being made manifest in what she terms scent marking, orSM, chiefly though not exclusively among males. Amongmandrills that might otherwise compete, a fresh kill may be shared.Although the literature does not reflect this side point, such behaviorcould be not so much a function of a sociology of food sharing asconsistent with the relative independence with which terrestrialranging/foraging activity is undertaken in the normal course of feedingamong baboons. (1974). (1995). Phillips, R. 161). Wickings, E. To begin with, thedistribution of adult males to adult females was about one to 14. References Bercovitch, F. Almost human: a journey into the world of baboons.New York: Random House. Among mandrills in captivity, same-sex grooming predominates(Stammbach, 1987). M., Collins, D. The physical reproductiveabilities and performance of males confined to the periphery of the troopappear to decline as well, although on a gradual basis (Dixson, Bossi, &Wickings, 1993). 118)that the aggression shown by male baboons is inversely related to thelength of time spent in a group; in fact, newcomers were the most dominant(aggressive) males in the troop, followed by short-term residents, with thelong-term residents last (in terms of dominance). Especially threatened in this regard are the mandrill and drill, whichare elusive and therefore problematic to study. B. The activity of grooming is, however, also evidence ofaffinitive socialization, which occurs among both mandrills and savannababoons. W. Mueller Graf, C. In some cases, this SM was followed by rubbing the breastas well. Primate Societies, pp. A. Kudo, H., & Mitani, M. C., Barnes, R. Napier (1974,p. E. American Journal of Primatology, 4 , 297-313. P., Barrow, B.

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