Variance in male reproductive success in single and multi-male red howler (Alouatta seniculus) troops was examined through paternity exclusion using genetic markers. Coalitions of relatives were compared to coalitions of non-relatives with regard to duration, stability, and number of offspring expected for each participant based on paternity exclusion results. No evidence of paternity by males living outside of the troop was found. In multi-male troops, only the dominant male was found to father offspring conceived during his tenure. This was the only male observed to mount females. In both troops examined genetically wherein a status change had occurred, paternity changed with status. Coalitions of relatives and potential relatives lasted significantly longer than coalitions of non-relatives, and experienced fewer status changes between males. A direct advantage of forming a coalition for both dominant and subordinate red howler males resulted from the superior competitive ability of coalitions over single males in establishing and maintaining tenure in the limited number of female troops in the population. Although the mating success of a subordinate male in a coalition appears to be extremely low, his chances of achieving membership in a bisexual troop as a single male may be even lower. Since the majority of troops in the population during any given year were multi-male or age-graded male (85%-96%), the number of troops that a single male would be able to invade unaided was very small. A subordinate male may be able to assume dominance, particularly if he is a subadult. In coalitions in which males are related, the subordinate male will further benefit from inclusive fitness. A dominant male that forms a coalition with a relative can expect to have longer reproductive tenure than in a troop with a non-relative, thereby producing more offspring and directly enhancing his fitness. Anecdotal evidence suggests that males choose relatives over non-relatives as coalition partners when possible.

Abstract. -The relationship between social structure and partitioning of genetic variance was examined in two red howler monkey populations (Wand G) in Venezuela, one of which (G) was undergoing rapid growth through colonization by new troops. Rates and patterns of gene flow had been determined through radiotelemetry and direct observation data on solitary migrants, and 10 years of troop censusing. Standard electrophoresis techniques were used to examine 29 loci in blood samples taken from 137 of the study animals. Analysis of genetic variance demonstrated: (I) a significantly high level of genetic variation among troops within populations (FST = 0.225 for Wand 0.142 for G), and (2) a significant excess of heterozygosity within troops relative to expected (F ,s = -0.136 for Wand -0.064 for G), despite relatively high levels of observed and inferred inbreeding in W. Differences between the populations in F S T values conformed to those predicted based on differences in colonization rate. Comparison of partitioning of genetic variance among different genealogical subsets of troops demonstrated that the pattern of genetic differentiation observed among troops within populations was promoted by an essentially single-male harem breeding structure, a very low rate of random exchange of breeding males among troops, and a high degree of relatedness among troop females. Between-troop genetic differentiation (F sT ) was thereby increased relative to that expected from other types of social organization, while the correlation between uniting gametes within troops (F,s) was decreased. Genetic differentiation between populations (2%) corresponded to that predicted from migration rates. Such a mosaic of genetic variation, combined with differences in reproductive success observed among troops and a high troop failure rate, create conditions in which interdemic selection could result in more rapid spread of advantageous gene combinations than would be expected in a panmictic population, particularly in a colonizing situation in which the founder population is small.

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Venezuelan red howler monkeys in a semideciduous habitat typically survive injuries and disabilities. Intraspecific physical aggression was the most frequently observed and inferred cause of injury. Thirty-eight percent of 119 howlers of all ages examined during capture had scars or other evidence of "damage." Overall, the sexes did not differ significantly in total number of injuries or number of individuals classified as "damaged." The incidence of injury was not independent of age-sex class. The subadult male class had the highest percentage of "damaged" individuals. However, when total injuries per age class were examined for each sex separately, only females showed a pattern that was significantly different than expected based on age (i.e., exposure to injury factors). Subadult females experienced more injuries than expected, whereas adult females had fewer injuries. Troop status (resident troop, natal troop, or extratroop) was not significantly related to the number of injuries in adult and subadult males. This finding was not surprising, because adult and subadult males of all status classes are involved in aggression related to breeding competition. Extratroop females had more injuries than expected, and natal females had the fewest injuries. The higher incidence of injuries on subadult females and extratroop females is consistent with aggression-mediated emigration of some females and observed resistance to female immigration by resident females. Overall, 74% of injuries were located on anterior-ventral portions of the body, consistent with the face-to-face fighting observed in howlers. Sociobiological costs of aggression in red howlers are difficult to assess because many howlers, despite frequent and severe injury, subsequently survive and reproduce. We recommend caution in inferring mortality from injuries, especially when social mobility and emigration out of the study area are common.