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Sexual Interference in Stumptail Macaques, and Where Qtpropriate, Data on Their Affiliatioa and Dominance

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Sexual Interference in Stumptail Macaques, and Where Qtpropriate, Data on Their Affiliatioa and Dominance SEXUAL INTERFERENCE IN STUMPTAIL MACAQUES (MACACA ARCTOIDES) IS IT RETURN-BENEFIT SPITE? ALYN ROBERT BRERETON Thesis submitted for the degree of Doctor of Philosophy University of Stiriing March 1989 ACKNOWLEDGEMENTS The mere fact that this thesis is coDq>lete is uttmishiiig. Why this is, I leave to legend and foiklore. But because it is, I alone am not entirely responsible. Many people and several organisations have unselfishly contributed, and I wim to acknowledge their help. I would, therefore, like to express my sincerest and deepest dunks: For financial s t^ o r t to: the Harris and Eliza Kempner Educational Fund the Hatde M. Strong Foundation Sigma K , Ihe Sdratific Research Society For techiicai assistance to: Dr. William McGrew Kristin Phillips Aiison Hannah Mario Caba Antonio Jauregui Dr. Ranald MacDonald Bnioe Sutheriand Dr. Arnold Chamove Ernesto Rodriquez Simon Booth KenDodcery Fm-overall encouragement to: Dr. William McGrew Krisdn Phillips Barbara Balen Janice Chu Debbie Bird Alison Hannah Dr. Liz Williamson FranDolins Eluned Price Dr. Arnold Chamovc Dr. Andrew Jahoda A m uFeisner For friendship to: Kristin Phillips Barbara Balen Dr. William McGrew Alison Hannah Dr. Liz Mlliamson Mark) Caba FranDolins Eluned Mce HofirioCanillo UUses Aguiloa Antonio Jauregui Dr. Arnold Chamovc ABSTRACT The phenomeiioa of intervening with other’s matings, or sexual interference, was studied in a fiee-tanging group of stumpcail macaques (Macaco arctoUUs). This thesis suggests that its function may be immediately spiteful, but also ultimately beneHcial reproductively. Preliminary evidence in support of this contention is presented, and speculations on related questions ate offered. A proposed evolutionary scenario of the stunqxail macaque mating pattern is «1«« outlined. CONTENTS F ite DEDICATION QUOTATION ACKNOWLEDGEMENTS ABSTRACT nC U R E S TABLES Chapter INTRODUCTION Ratioaile Objectives duquerSumnaiies STUDIES OF SEXUAL INTERFERENCE Descriptive Overview Invertebmes Iniectt Veitebntes Fish Aonhibians Rqxfles B ins Msmonls Rodentt Aitiodictids C m ivores Pinnipeds Elephants PrinutBS Prosimians New World Monkeys Old World Monkeys Apes Humans Specific Studies and Rndings Insects Fish Amphibians Pinnipeds Fimates Summary RESEARCH SETTING Taxomiiqr of StunnaU Macaques (MacaeamtoUia) Status Social devious Studies of Social Bduviour Wild Captive Study Site Locadon F lon and Fauna O im ie Sub|ect'sHistOfy METHODS Subjects ^ip iratu s Procedures Samiding Rotocols Analysis Measures Statistics THEORETICAL FUNDAMENTALS OF SEXUAL SELECTION AND SOaOBIOLOGY Sexual Sdection Sociotmlogy SOaO^EXUAL BE3UVIOUR Macaca Review M.SUemu-Sytfama Group Social Aoributes Sexual Attributes M. Fasdcularis Qmap Social Attributes Sexual Attributes M. Sinica Group Sodai Attributes Sexual Attributes M.Arctoides Gn>«q> Social Atufixites Sexual Attributes Description of Stumptail Mating, Copulatoiy Tie, and Genitalia Morphology Mating Desciqttion Copuuitoiy'ne Getiitalia Moiphdogy Male Female Result! OiDup Individual CoimMrisoiu and Discutiioa Gromi Individual SEXUAL INTERFERENCE 117 Distributioa of Sexual Interfoenoe Among Mmates 117 Detoripdoa of Sexual Imerfeieiioe Among Stunqxails Resultt Group Individual Coomaiisoas and Discussion Group Individual SEXUAL INTERFERENCE: AS SPITE? Ptevious Ifypotheses Effectt of Stress on Rqxoduction Resultt O nM adnp On Individuals Between Individuals Reproductive Data IBstofical Study Period The Spite Hypodwsis Predictive Tests (1) Resultt (2) Resultt (3) Resultt (4) Resultt (5) Rqm ductive Data (Q Rqm ductive Data 0 Rqxoductive Data Discussion Evidence Resultt Predictive Test 1 nedictiveTett2 Predictive Test 3 I¥edictiveTest4 Reproductive Data nedicdveTestS I¥edictiveTest6 RedicdveTest? I or Further Tests Stumptails Other Macaques SEXUAL INTERFERENCE: AS RETURN- BENEFIT SPITE? C oom int-Im Reproductive Siruegy Alieniaiive Rnroductive Tactics Stumptail Tactics Results General Specific Reproductive Data The Return-Benefit Spite Hypothesis IVedictive Tests (1) Results Q) Results O ) Results (4) Reproductive Data Oiscussk» Evidence Results Predictive Test 1 nedictiveTest2 Predictive Test 3 RqmductiveData RedictiveTest4 Reproductive Value S|rite with a Reiuro-Benefii? 221 10. DISCUSSION AND CONCLUSIONS Discussion Sexual Interference Madng S etiw Interference and Mating Evolutionaiy Stages (rfStumpiail Mating? Snuctural Stage 1 S u ^ 2 Behavioural Stages Staged Conclusions APPENDIX Ternis and Definitioas of Bchavknir by Category REFERENCES 245 FIGURES Figure Page 3.1 Taxonomy of dw genus Miocaca. 36 3.2 Oeogn^ihic distribution Macaco in Asia. 38 3.3 Geogrq>hic locatioo of Tananrillo in Lake Caiemaco, state of Vencniz, Mexico. 41 3.4 Elevations in the San Martin Tuxtla mountain range, Veracruz, Mexico. 42 3.3 location of Tanatqrillo within Lake Catemaco, Veraouz, Mexico. 42 3.6 Dimensions of T a n a t^ o within Caiemaco, Veracruz, Mexico. 44 3.7 Tanatqtillo from a distance 0>hoio centre) showing the Tuxda range in the backgroiimd. 45 3.8 The two portions (rfTaiuuqtillo showing an outline of its canopy. 45 3.9 Another view of TanatqriUo's canopy. 46 3.10 Partial view of Tanuqnllo looking soudiwest towairi the town of ratem im 46 3.11 Partial view of Tanatqtillo looking west. 47 3.12 Partial view of Taiuuqrillo looking east toward die more open area ^ the Ivger of the two islands. 47 3.13 Provisioning of bananas. 50 3.14 Provisiai of bananas contrived by Pancho (centre right) and McGregor (centre left). 30 3.15 Provision of cmn. 51 3.16 Supplemental feeding by stunqitails on nannlv^etation. 51 3.17 Airival of a tourist boat to the island. 53 3.18 StumptailAourist interaction. 53 4.1 GnMuconqMsiiioaoiiTa HMlriline fimn N6void)er. 19^3 diroagh May, 1984. 57 5.1 A hypothetical I 76 6.1 Genitalia n io n )h o k ^ of nude Afoooea arcÊoides. 94 6.2 Genitalia motphotogy of fenude Afacoca araotdes. 6.3 Matings by month. 6.4 Matings by hour of the day. 6.3 Mating behaviour averages. 6.6 Male mating distribution. 6.7 Female mating distribution. 6.8 Coiielation scattergram tA afiBUation duration widi mating fiequency. 7.1 McGregor mating with Rosy. 7.2 McGic|orwithafrown-n>und-mouth expression which signals ejaculation. 122 7.3 McGregor tied, sitting back, and threatening an intnider, wMle interference (mild in this case) continues by an individual in the for^round. 123 7.4 Rosy and McGregor remain tied McGregor continues threatening. 7.3 Tying and threatening continue. 7.6 McGregor stands and continues threatening. 7.7 Percent of matings receiving sexual intoference. 7.8 Perçoit of individuals interfering 1^ sex. 7.9 Percentages of matings by numbered intruders. 7.10 Percent of sexual interferoice by sex. 7.11 Fercem of Mxiial intafiEreiioe by group. 132 7.12 Peicentof lexml interfeience hy nMtrilim» 133 8.1 H k role of honnones in female (niinate ovulation. ISO 8.2 Ovuladoo feihire in female mannoiets (CaWtMxJaccim) produced by hotmonal stqtpfession and induced by lodal stress. 1S3 8.3 Mating behaviour with and without sexual imefference. 1S6 8.4 Mean tie dotations of first and second halves of serial matings receiving sexual inteiferaice. 1S7 8.S Variance fium overall mean tie duration by number intruders. 1S8 8.6 Mean number of pre-ejaculatciy mounts for matings with and without sexual interference. 160 8.7 Pierre's, Flaca's, Bruja's, and Rosy's mating distributions widi males. 164 8.8 Average number intruders per serial mating by first and second halves for Pancho and McGregor. 168 8.9 Average number of inttuden per serial mating by first and second halves for neiie, Flaca, Fanny. Luda, Btuja, and Rosy. 169 8.10 Variation among males in tie duration: overall, with, and without sexual interference. 171 8.11 Vuiation among females in tie duration: overall, with, and without sexual interfeience. 172 8.12 Average number of inmiders per iruuing for males. 174 8.13 Average number of intruders per iruuing for females. 17S 8.14 Average interval betweoi matings for males. 177 8.1S Average interval between matings for females. 178 8.16 Fnnale age at first riving birth for the Tanaxpillo stumptuls over a IS year period. 180 8.17 Numbaof live biitlu ainoiM the feoales Jarodu. Margarita. Flaca, BkmiUe. and Flor. 181 8.18 Mean im atiith intervals for the Tanajqrillo fanales over a IS year poiod. 183 8.19 H e duration o{ possible oonontioa mating compared with mean tie durauon for matings overall. 185 8.20 Amount of seaual interference determined by total intruder oomlnned score lor possible cono^)6on mating congiaied with the mean combined score for mating overall 186 8.21 The beta male. McGregor. 190 8.22 The alpha male. Pancho. 191 9.1 Peroouage alternative tactic matings observed out (rf the total of 113 matings. 204 9.2 P ercoita« of aliemative tactic matings observed^out of the 19 matings recorded without sexual interference. 205 9.3 Percentage of the 30 recorded alternative tactic mitin|s recoving sexual interfermce compiued with dw percentage receiving interlermce among the 83 other matings. 206 9.4 Average amount sexual interference per mating for the 30 alternative tactic matings opposed to the 83 odwrs. 207 9.5 Average amounttrf sexual interference per mating for both the 19 female leave and 11 male mde-and-hurry matings, 209 9.6 Mean tie duration for the 19 female leave and 11 male hide-and-huny matings. 210 9.7 Dififiteiices among females and males in the average amount of sexual interference received per mating for die two Qtpes of altemadve tactic matings. 211 9.8 Differences among females and males in mean de duradon for the two types of alternative tactic matings. 213 9.9 Avenge number months between births for the 3 females who practised alternative tactic matings compared with the 7 who did not 215 TABLES Table Page 2.1 Sumnaiy of texutl interference among animal ipecies. 5 3.1 Composition (rf the two groim of stumptail macaquM during August anoNovember. 1974. 49 4.1 Nundier of subjects on TanajqxUo by subgnxq) age and sex. 55 4.2 Otoiq> composition on TanaxpiUo individual s u lg ^ from November. 1983 tfa ro i^ May.
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