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Papio Anubis) and Vervet Monkeys (Chlorocebus Pygerythrus

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Papio Anubis) and Vervet Monkeys (Chlorocebus Pygerythrus MONKEYING AROUND: COMPARING PLAY BEHAVIOR IN RELATION TO PREDATION RISK IN SYMPATRIC OLIVE BABOONS (PAPIO ANUBIS) AND VERVET MONKEYS (CHLOROCEBUS PYGERYTHRUS) Danielle Steinberg Abstract: Predation has long been considered a strong selective pressure on primate behavior, including foraging and feeding. However, its effects on play behavior, a crucial aspect of primate life, are unknown. Feeding is necessary for survival, but play is imperative for later reproductive success. Olive baboons (Papio anubis) and vervet monkeys (Chlorocebus pygerythrus) are often sympatric but baboons are larger and more aggressive, suggesting that they may be less vulnerable to predation and thus may have longer or more frequent play bouts. Similarly, within species, male primates, being more aggressive than female primates, may be less vulnerable to predation and so may play more than females. I observed play behavior of baboons and vervets in Nyungwe and Akagera National Parks, Rwanda, to test these predictions. I used all-occurrence sampling to record frequencies and durations of play bouts and analyzed the data with nonparametric statistical tests. There were significant differences in play bout frequencies but not durations both between the species and between males and females within species. Future research should explore whether life history differences also contribute to differences in play behavior. Keywords: Vervet | olive baboon | play | predation Introduction the habitats proportionally to their availability and they spent less time in food-rich areas Primates and their predators have been in an than expected due to the higher risk of evolutionary arms race for over 70 million predation in those areas. In a similar study, years (Isbell, 2009). The effects of predation Coleman and Hill (2014) found that samango as a strong selective pressure on primate monkeys (Cercopithecus mitis erythrarchus) behavior have been studied in numerous decide on habitat usage based on predation contexts. Ecological tradeoffs lie at the root of risk rather than on resource availability. predation as a selective pressure, specifically in Ultimately, predation may lead to trade-offs in regards to feeding and foraging behaviors and foraging, habitat use, or another common habitat use (Cowlishaw 1997; Lima 1998; Hart behaviors in primates (Lima 1998; Hart 2007). 2007; Coleman and Hill 2013). For instance, Though predation has obvious effects in a study conducted by Cowlishaw (1997), on the daily lives of all primates, certain chacma baboons (Papio ursinus) were characteristics may make some primates more observed under different conditions of prone to predation than others. Smaller- predation risk and food availability in a given bodied primates are more susceptible to habitat. He found that the baboons did not use predator attacks than are larger-bodied © D. L. Steinberg 2017. Originally published in Explorations: The UC Davis Undergraduate Research Journal, Vol. 19 (2017). http://Explorations.UCDavis.edu © The Regents of the University of California. primates (Isbell 1994; Enstam 2007). For this widespread theory on the functionality of play reason, different primate species tend to have is honing social skills that can be utilized different predator avoidance or deterrent throughout the animal’s life (Chalmers 1987; strategies. Shultz et al. (2004) noticed that Lewis 2000; Fairbanks 1993). Play often terrestrial primates are more likely to live in necessitates cooperation and communication large groups or to be large in body size as a between partners which inherently works to predator deterrence strategy. One hypothesis increase social competency. Lewis (2000) regarding extreme sexual dimorphism in some noted that primate social play was positively species is that large males act as predator correlated with neocortex ratio. She attributed deterrents, implying that large body size does, this to the fact that social play requires a in fact, reduce risk of predation (Leutenegger number of complex cognitive function such as and Kelly 1977). In addition to large body size, predicting behaviors and reacting alarm calls are another popular predator appropriately which, in turn, helps the animals deterrent (Enstam 2007; Isbell and Bidner develop skills useful in their adult lives. 2016). In a study by Enstam and Isbell (2002), Fairbanks (1993) noted that even the motor predation responses were compared between skills acquired through play can aid in primate vervet monkeys (Chlorocebus pygerythrus) reproductive success. She argues that male and their slightly larger relatives, patas primates develop fighting skills through play monkeys (Erythrocebus patas). They found and that fighting ability in male that vervet monkeys tend to alarm call and flee cercopithecines often precipitates greater while the larger, more aggressive patas reproductive success. Social competency is monkeys would sometimes engage in “active even more crucial to reproductive success defense” (Isbell and Enstam 2002). Like patas because successfully integrating into a new monkeys, red colobus (Piliocolobus group post-dispersal requires appropriate tephrosceles) males have also been observed to social skills (Platt et al. 2016). Ultimately, use aggression as a predator deterrent playing during the juvenile period helps build (Struhsaker and Leakey 1990; Boesch 1994; the skills necessary for later acceptance in Mitani and Watts 1999). In three separate future social circumstances including dispersal studies, researchers noticed that chimpanzees and, later, courting behavior. attacked younger red colobus and avoided Even though play eventually produces adult males due to their aggressive responses benefits, there are immediate costs. Play is a (Struhsaker Leakey 1990; Boesch 1994; Mitani very risky behavior because it puts individuals Watts 1999). in precarious situations and leaves them Just as all primates are subject to susceptible to predation (Struhsaker and predation risk, all primates play. Play is Leakey 1990; Lima 1998; Fagen 1993). During defined by Burghardt (2005) as satisfying play, individuals are not completely aware of several conditions including arising their surroundings and are prone to falls. A spontaneously, having no immediate function, number of studies involving lethal falls cited by and occurring in a stable environment. While Fagen (1993) indicate the high risk that often many primatologists agree that the benefits of correlates with play. Furthermore, play play occur much later than when the act itself reduces the capacity for vigilance as attention takes place, the specific benefits are still is directed toward the object of play rather debated (Chalmers 1987; Pellis 1988; than surroundings. For instance, in a study of Struhsaker and Leakey 1990; Lima 1998; yellow baboon (P. cynocephalus) predatory Lewis 2000; Fagen 1993; Fairbanks 1993; behaviors, Hausfater (1976) noted that the Maestripieri and Ross 2004). Developing baboons preyed upon mostly younger vervets good motor skills is one of the primary who were in play groups away from the more hypothesized functions of play (Fagen 1993; vigilant adults. In addition to reducing Maestripieri and Ross 2004). A second vigilance, play makes individuals more UC Davis | EXPLORATIONS: THE UC DAVIS UNDERGRADUATE RESEARCH JOURNAL Vol. 19 (2017) Steinberg p.2 conspicuous to their predators. In its most previously, both body size and aggression have simple yet justifiable form, increasing activity been shown to be negatively correlated with leads to an increased risk of predation (Lima predation risk so baboons should therefore 1998). Struhsaker and Leakey (1990) found have a lower risk of predation than vervet that crowned hawk-eagles (Stephanoaetus monkeys do in the same general environment. coronatus) did not randomly select primate Here I test the prediction that risk of prey, but rather targeted adult males in four predation affects olive baboon and vervet primate species (except red colobus as noted monkey play behavior differently. Specifically, above) because they tended to be more I predicted that baboons would have both demonstrative, give more alarm calls, and be longer and more frequent play bouts compared less cautious overall which made them very to vervets due to lower risk of predation conspicuous, easy targets. Similarly, play because of their larger body sizes and more reduces vigilance and increases noticeability aggressive nature. To examine the relationship making the players easy targets for predation. between predation risk and body size and Evolutionarily speaking, risky behaviors aggressiveness further, I also predicted that should not be performed unless they have female baboons would play more often or for benefits that outweigh the costs. This longer than female vervets, male baboons necessitates then that play does in fact lead to would play more often or for longer than male benefits such as those stated above in order for vervets, and within each species, males would the tradeoff to be worth it evolutionarily. play more often or for longer than females. While play involves risk of predation, the risks themselves vary across species and habitats (Enstam 2007). In order to account Methods for these variables, I selected vervet monkeys and olive baboons, two species very similar in Study sites habitat and that share common predators, to I collected data from June 14 to June examine how predation may affect play 30, 2016 in two of Rwanda’s national parks: behavior. Vervets and olive baboons both
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