R. Boonratana et al.: Mating Relationships | THE DEVELOPMENT OF MATING RELATIONSHIPS IN A SMALL GROUP OF MA- CACA MULATTA. R. BOONRATANA and C.J. EDWIN. Introduction ‘ Sexual behaviour of the Macaca mulatta has been rider extensive observations since the fa~ mous studies by CARPENTER (1942a and b). The studies by KAUFMAN (1965) and DRICKA- MER (1974) have formed the data base with which other investigators of rhesus sexual behaviour have often made comparisons. Much of the conceptual framework leading to the understanding of rhesus sexual behaviour has been reviewed by AGAR and MITCHELL (1975). LINDBURG (1971 and 1975), MITCHELL (1979) and FEDIGAN (1983). ‘The object of this paper is to report the development of mating relationships i ina small, natu- rally occuring group of Macaca mulatta. Against the background of the genealogical structure, the mating relationship that existed at the beginning of the study and the new ones that appeared in the course of it are analysed and interpreted in light of known theory. The hypothesis presented are offered for validation in larger and ecologically diverse rhesus groups. Methods Focal animal sampling (ALTMANN, 1974) has been used with observation being made with a frequency of 15 seconds in 300 second blocks, every half hour from sunrise to sunset. The obser- vation began in February 1985 and at the time of reporting are still continuing. Occurences of be- haviour were recorded for a set of predetermined elemental patterns. These records were supple- mented by notes made during the observation of the group. The group members were in all ages (age-sex structure being classified following SOLTHWICK et al., 1965). Barring the two infants (less than one yéar od) the focal animals were sclected from among the other nine members of the group. The total number of observations till end of January 1986 were 13200 samples. The group is usually found in the wooded region West of Chandigarh city (India), According to reports the group formed some 7-9 years ago (personal observation, EDWIN, C.J.). The first members were one adult male and two adult females. Observations 1, The Group ‘The size of the group has varied during the course of observations, being 9 during the period October-December 1984 with two binis in Api and May 1985 the group had increased to 11 members. The loss of two adult males in the period December 1985 to May 1986 was offset by 2 more births in May and June 1986, Hence, the group retained the size of 11 members. Figure | represents the composition and kin structure of the group as in September 1985. The group is composed of three elements. There arc two natal groups and the third element is described as 'others'which comprises the alpha male (Kong) and three subordinate males. Primate Report 15, December 1986 21nana et al.: Mating Relationships joonrat R. Be P 4 (pouxtsuoaun x9s) g¥gi ‘9 auNE UO ,UaTuEG, 03 yIZTq eALS Apne , (8Tew) “yest f0z ABW UO ,32BWeS, 07 YIITG aAes sau0H 2 $861 19quad0qg us—s ION .. : i oest kew UE pera . \ $861 1eqwaadas ur sy (eTew 3tnpe-qns) (© ew atnpy) (atew 3tnpy) “ee (1A) 0Z08 (A) aaD4ve (AL) Lranva (9cewss syau sj . (sak stg apeusg) (etew sya p) (sak sz ereN) (S24 s*¢ ateH) (x) VeNIN (IITA) YBONTHS (x) Ila (x1) ATIOr (IIA) o£oL ya (eT euey acnpy) oy v- + (9pRmaz arnpy) Bs oT een CTE) BEHOW Pose oC) kane (#1ew 3tnpy) (1) 9Nox TeNpyAypuy Jo wey pu, UOFaedTzysSe{> xos-o¥e Burmoys dnos8 30 osngonsas dyysury 21 “Sra December 1986 Primate Report 15, 22ips ionshiy Mating Relati E Boonratdth etal. [v102 jo wee. 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These are: 3 i i) Friendly or cooperative behaviours i) Antagonistic or competitive behaviour iii) Neutral behaviours. | | I, Table 1 represents the proportion of theie types of behaviours expressed as.a fraction of the total social interactions seen in each month between the period February 1985 to January 1986.As is evident that the large proportion of interactions are friendly and pooling in the dyadic and sub- 2roup interactions (more than 2 participants). We find that friendly interactions are usually above 0.9. Itis expected that the origins of the processes which lead to the formation of mating rclation- ships would lie in the manifestation of cooperative social behaviours. This is accentuated by the fact that social behaviours (except in March 1985) form the larger proportion of otal behaviours. ' “Table 2: Vartation in principle behavioural categories Within Between Between Among . own natal = two natal natal groups others group groups and others 63.51 10.81 Aggression 2.7 63.51 Resting 43.41 23.97 32.00 0.84 Feeding 21.73 47.63 13.93 16.71 Playing 23.38 60.39 16.23 - Sexual - 34.48 65.52 - Mobility 82.01 - 17.99 - ‘The patterns of the social interactions seen for principal behavioural categories presented in Table 2 indicate that there is a variation among them. Proportionately aggression is highest be- tween members of the natal groups and others. This was principally a consequence of the aggres- sion with the subordinate males. Resting and moving together were more often exhibited by members of a natal group within themselves. Feeding and play behaviours are areas of coopera~_ tion between the two natal groups and sexual activity was for the entite observation highest be- tween members of the natal groups and others (principally Kone). Mating :ithin natal eroups was not scen. Based on these observations we fee! that the pracesses of the formation of mating relationships would lie within social groupings engaged in cooperative behaviours. ‘ocial Groupings a Itis assumed that selective processes are involved in the exhibition of subgroups ranging in size from 2 to 10, of, it may be treated as the exclusion of menthers ranging from 9 to 1. : Table 3 shows the expected and observed number of combinations of subgroups, sizes rang- Primate Report 15, December 1986 - i 24 | | SeR, Boonratana et al.: Mating Relations ips Table 3: Expected and observed number of sub-group combinations Sub-group = . % Expected" i . Observed “size 4 . “bohbination +" .combination ? ' uM ia ; 2 + ; “e s E 2 Fey 10 6 ‘ 2 7 7 4 6 8 ‘ = “ se} 9 % ' ~ 1 A | es eee Nil “2, ps. 005; df ing from 2 to 10. The observed number of combinations are highly significantly less than (using ) the expected which indicates that association is not random but highly selective. ‘The participation of members of the group in various subgroups varies. The modal trend is for the attainment of complete participation (frequency 1.00) as the subgroup size rises. Table 4 shows the frequency of parti different members. Table 4: Participation levels of various group members in” sub-groups » of different sizes 5 M GROUP SIZES Members | ~ 2 3 4 5 6 7 8 9 Kong 0.267 0.381 0.643 0.500 0.857 0.833 1.009 1.900 Judy 0.233 0.476 0.429 0,400 0.714 1.000 1.006 1.000 Momee 0.233 0.190 0.357 0.300 0.714 0.833 1.900 1.000 Bandit 0.133 0.048 = | - - - - - Jaycee 0.033 = = - - - - - Bozo 0.200 0.190 0.071 0.100 = 0.333 0.500 1.000 Toto” m 0.200 0.381 0.357 0.700 0.571 0.833 1.000 1,000 Swinger 0.233 0.333 0.714 1.000 0.857 0.833 1.000 1.000 Jolly 0.267 0.572 0.714 0.900 1.000 1.000 1.099 1.900 Ninja 0.133 0.238 0.357 0.700 0.714 0.667 1.900 1.000 Fifi 0.067 0.190 0.557.0.400 0.571 0.667 0.500 1.000 Primate Report 15, December'1986 ipation in subgroups of various sizes (ranging from 2.109) byR. Boonratana étal.: Mating Relationships “ing uofzedposzued yo Aduanbaus 9.9 Figure 24: Participation by group members in sub groupings (size=sn = 2 to 9) ert i Primate Report 15, December 1986* R. Boonratana et al.: Mating Relaionships From the graph (Figure 2) which is the juxtaposition of all the individual participation frequencies (excepting Ninja and Fifi which are omitted for sake of simplicity), we see that there are three ob- servable clusters which can be identified in decreasing onerof participation ar Lolly and Swinger’ _ IL Kong, Judy, Momee, Toto (Fifi and Ninja) III Bandit, Jaycee and Bozo : : ‘The graph shows the relative social exclusion of the three subordinate males from the configura- tion of friendly relationships and by extension of the argument, exclusions from the mating struc- ture. _ ‘s ass oe STR . ‘The data cover two mating seasons, in 1984 and 1985. es J Asis evident from Figure 3 that at the close of 1984, Kong was mating with both the adult females, Judy and Momee. However in the period prior to the onset of the mating season of 1985 Kong was seen exclusively with Judy."Although Kong and Momee were seen for prolonged periods to- gether no mounting was Seen between them. Momee initiated consort relationships with Toto (Judy's maturing male offspring). With the extended mating season into the early part of 1986 Momee and Toto were always seen together and in reproductive relationships. [tis our submission that the offspring born to Momee in May 1986 was fathered by Toto. ‘The data on social relations obtained in the course of 1985 shows prolonged yearlong contact of Swinger and Jolly. Although these two were always together there was no instance of any se- xual mounting. Play mounting however did exist. It must be noted that both were sexually imma- ture. Apparently this relationship may be described as juvenile association but it would not be entirely unexpected that in later part of the mating season of 1986 this relationship acquires a se- * xual character. Figure 3: Mating relations observed in October-December 1984 and November-December 1985, TOTO" Joly FIFL SWINGER NINJA BANDIT JAYCEE BOZO —— Mating relationships seen in October-December 1984 - Mating relationships seen in November-December 1985 ---:- Seen together in November-December 1985 but no sexual mounting. Primate Report 15, December:1986 : 27R. Boonratana et al,: Mating Relationships ei Discussion Thisis a case study ona newly formed group of the Macaca mulatta. The group most of the groups reported in literature, however, this fact permits close observations leading to the formation of hypotheses. This study rests on the presumption that mating rélationsips in any social group are a part of; ; the entire gamut of cooperative behaviours existing within the members. Further, that howso- ever, small a group may be, social mechanisms exist to prevent surplus males from becoming part of the breeding structure, although these males for all intents and purposes are a part of the social group. The breeding structure it is proposed is a separate entity from the'social groups. This breeding structure is the consequence of the choices exercised for mates by both males and fema-", les. The data of this study support the hypotheses of LINDBURG (1975) that female choice plays an important role in the formation of breedjng structure. In this study Momee (adult female) op- ted for a maturing male to the exclusion of three other mature subo1 te males. Was this mani- festation motivated by a positive selection for dominance? This would be the initial conclusion but the data are insufficient to be able to speak of it definitively. Finally the results of this study show that even in very small sized groups of the ihean were choice of mate is numerically small there is inhibition of mating within natal groups, & position earlier established by the studies of SADE (1968), DUGGLEBY (1977) and MISSAKIAN- QUINN and VARLEY (1977). Thus the development of mating relationships in the rhesus is a consequence of preferences by males and females in a socially established breeding structure . which serves to increase genetic variability without appare:it arm to the whole social group. As far as the focal group is concerned, observations fall within the existing theoretical framework of rhesus sexual behaviour. Thereby this makes for the acceptance of many small sized rhesus groups as social units. Acknowledgements The authors are grateful to the Chairman, Anthropology Department, Panjab University, Chandigarh for facilities extended. Panjab University Chandigargh (India) is gratefully acknow- Jedged for financial support to one of us (C.J. EDWIN). The authors also gratefully acknowledge the assistance reccived from Mr. K. BOSE and Mr. Chain SINGH. tiie References AGAR, M.E. and MITCHELL, G.: Behaviour of free ranging adults: a review, In Bourne, G. (ed.). The Rhesus Monkey, Vol. 1, Chapt. 8, pp. 323-342, Academic Press, New York. (1975). ALTMANN, J.: Observational Study of Behaviour: sampling methods. Behaviour, (1974) XLIX, 227-267. CARPENTER, C.R.: Sexual behaviour of free-ranging Rhesus Monkeys (Macaca mulatta) 1 Specimens. an Procedures and Behavioural Characteristics of Estrus.J. Comp. Psychol.,(1942a), 33: 113-142, 28 ‘ Primate Report 15, December 1986©" R. Boonratana et al.: Mating Relationships CARPENTER, C.R.: Sexual behaviour of free-ranging Rhesus Monkeys (Macaca mulatta), II: Periodicity of Estrus, Homosexual, Auloerotic and Non Conformist Behaviour.J.Comp. Psy- chol., (1942b), 33: 143-162. DRICKAMER, L.C.: A iemyear Summary of reproductive Data for Free-ranging Macaca mu- Jatta.Folia Primat. (1974), 21: 61-80. DUGGLEBY, C.R.: Inbreeding in Macaca muilatta from Cayo Santiago. Paper presented at American Society of Primatologists Meeting, Seattle, Washington,(1977) (cited from MIT- CHELL, G.,.1979). FEDIGAN, L.M. : Dominance and Reproductive Success in Primates. Yearbook of Phys. An- throp., (1983), 26: 91-129. a 2 : KAUEMANN, J.H.: ArTseyea Study of Mating Behaviour in Free-ranging Band of Rhesus Monkeys. Ecology, (1965), 46: 500-512. LINDBURG, D.G.: The Rhesus Monkey in North India: an Ecological and Behavioural Study. In Rosenblum, L.A. (ed), Primate Behaviour: Developments in Field and Laboratory Research, Vol. II, pp. 1-106, Academic Press, New York, (1971). LINDBURC ™ G.: Mate Selection in the Rhesus Monkey, Macaca mulatta. Amer. J. Phys. An- throp., (1975), 42: 315. MISSAKIAN-QUINN, E.A. and VARLEY, M.A.: Consort Activity in groups of Free-ranging Rhesus Monkeys on Cayo Santiago. Paper presented at the American Society of Primatologists Meeting, Seattle, Washington, (1977) (cited from MITCHELL, G., 1979). MITCHELL, G.: Behavioural Sex Differences in Non-human Primates, Van Nostrand Reinhold Co., New York, (1979). SADE, D.S.: Inhibition of Son-Mother Mating Among Free-ranging Rhesus Monkeys. Science and Psychoanalysis, (1968) 12: 18-32. SOUTHWICK, C.H., BEG, M.A. De Vore, I. (ed.), Primate Behviour: art, and Winston, New York, (1965). J SIDDIGUI, M.R.: Rhesus Monkeys in North India. fn Id Study of Monkeys and Apes, pp. 111-159. Holt, Rineh- Authors’ addresses: | Anthropology Department, Panjab University, Chandigarh-160014, INDIA. Primate Report 15, December 1986 29