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Female spotted hyenas (Crocuta crocuta) are larger and more aggressive than males, and their genitalia are heavily 'masculinized'. These odd traits in females pose unusual challenges for males during courtship and copulation. Here our goals were to describe and quantify the behavior patterns involved in courtship and copulation in Crocuta, and determine whether rates of affiliative behavior directed toward females by males vary with female age, social rank, or time to conception. We also inquired whether consort formation with a particular female was necessary for a male to sire that female's cubs. Behavioral observations and paternity data based on 12 microsatellite loci were collected over 11 years from free-ranging hyenas in Kenya. Several of the courtship displays exhibited by male hyenas differed from those found in other carnivores, and appeared to reflect intense motivational conflict between tendencies to approach and flee from females. Most male advances were either ignored by females or elicited aggression from females toward males. Rates of male affiliative behavior toward females peaked around the time of conception. Although males behaved similarly toward young and old females in the highest social rank category, males directed more affiliative behavior toward older than younger females that were mid- and low-ranking. Multiple short mounts usually preceded a long mount, but intromission and ejaculation appeared to occur only during long mounts. Female receptivity was indicated by inhibited aggression toward the male and assumption of a distinctive receptive stance. The only behavior indicative of female proceptivity was following of the male by the female in mating contexts. Some males who sired cubs formed consortships with females whereas others did not, suggesting that individual male hyenas may adopt alternative reproductive tactics to attract and acquire mates. Our results also suggest that low fertility may be an important cost of female virilization in this species. [ABSTRACT FROM AUTHOR]

It has long been recognized that vocal signals communicate information about the age, sex and affective state of callers. However, the mechanisms by which these types of information are communicated are less well understood. Here we investigated variation in the acoustic properties of the long-distance vocalizations, called 'whoops', emitted by free-living spotted hyenas, Crocuta crocuta. Specifically we investigated whether the fundamental frequency, length and rate of whoops provide information about the caller's age, sex and/or level of arousal. We determined the latter by contrasting whoops emitted spontaneously with whoops emitted during periods of social excitement, when callers typically also exhibited visual signals associated with heightened arousal. We found that the minimum fundamental frequency of a whoop provides reliable information about the caller's general age and, for adult callers, information about sex as well. The vast majority of adult male whoop bouts were emitted spontaneously, but juveniles and adult females produced many of their bouts during periods of social excitement. Although context did not significantly affect the whoop bouts of adult females, juvenile bouts emitted during social excitement had higher maximum fundamental frequencies, greater proportions of asymmetric whoop subtypes, and reduced inter-whoop intervals. By reducing the inter-whoop intervals of bouts, juvenile hyenas significantly increased the likelihood that conspecifics would respond to whoops by approaching the caller or its social companion. Peak fundamental frequency and the relative abundance of whoop subtypes did not appreciably affect response. We discuss the potential functions of whooping by juvenile and adult hyenas in light of these findings. [ABSTRACT FROM AUTHOR]