Why Do Nazca Boobies (Sula Granti) Perform NAV Behavior?
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Zoe von Gerlach Professor William H. Durham October 15th 2018 Why do Nazca Boobies (Sula granti) perform NAV behavior? Abstract This research project focuses on Nazca boobies and their Non-Parental Adult Visitor (NAV) behavior. Non-breeding adult Nazca boobies will go to a nest that is not their own to interact with the unattended chick whose parents are out foraging for food. This interaction can either be playful, aggressive or sexually abusive (Tarlow et al 2001: 1). The purpose of this research project was to examine three different hypotheses dealing with the reasons why NAVs perform this behavior, looking specifically at whether male adult Nazca boobies are more aggressive than females and the reasons behind this. Through the literary research conducted, it was concluded that females are actually much more aggressive than males. The different factors influencing the frequency of NAV behavior as an adult include siblicide and personal victimization as a chick. Although these factors influence NAV behavior, it is likely that high hormone levels at birth and young age have organizational effects that cause this phenomenon in grown Nazca boobies. Introduction Figure 1: Map highlighting Sula granti’s range of occurrence (IUCN Red List) There are three native booby species to the Galapagos. The Red-footed booby, the Blue- footed booby and the Nazca booby. The focus of this project is the Nazca booby. They exhibit two special behaviors that make them so fascinating for humans to study. Firstly, they practice obligate siblicide. Although the typical clutch size of a Nazca booby nest is 2 eggs, only one hatchling ever reaches adulthood. The reason the female lays two eggs is out of insurance, since their hatching success rates are only around 60%. However, if both eggs hatch, then the stronger sibling will always kill the weaker one (Ferree et al 2004: 1). The second behavior Nazca boobies exhibit is NAV behavior. This typically happens when the chick is between 20 and 80 days old, young enough to not be able to defend itself but old enough that its parents leave it alone while foraging for food. These NAV interactions take one of three forms, either the interaction is playful, aggressive or sexually abusive. If the interaction is playful, the adult NAV will bring pebbles or feathers to the chick’s nest and play with it, exhibiting parental behavior. The aggressive interaction often involves jabbing and biting the chick as well as other types of aggressive advances. The sexually abusive interaction is the least common of the three, and involves the adult NAV trying to copulate with the chick (Anderson et al 2004: 1). One effect of aggressive NAV behavior is that it can indirectly lead to a chick’s death. The wounds created by the NAV attract landbirds to feed blood from the chick. This weakens the chick, and often results in its death (Anderson et al 2004: 3). This paper will assess three hypotheses and examine published literature to evaluate and asses them. Hypotheses 1. Adult male Nazca boobies that were abandoned by their female partner take out their frustration on young chicks through aggressive behavior 2. To improve their own chances of finding a mate, NAVs injure young chicks to reduce the chick’s physical appeal once they are grown (reducing the attractiveness of the competition) 3. Female NAVs have a higher percentage than males of exhibiting playful behavior towards chicks. This is caused by a combination of motherly instinct and hormones, specifically androgen Method There was no personal research conducted for this paper. Instead, this paper is an in depth literature review of published peer-reviewed studies on the subject. All figures are taken from these studies. All photographs were taken personally. When we were in the Galapagos, chicks had already matured enough to defend themselves against NAVs, so sadly there are no personal photographs of this specific phenomenon to contribute. Hypothesis 1 Adult male Nazca boobies that were abandoned by their female partner take out their frustration on young chicks through aggressive behavior Findings Figure 2: Graph showing the number of females and males committing aggressive NAV events (Tarlow et al 2001: 4) This hypothesis is not correct. Firstly, as figure 2 shows, female NAVs are actually more aggressive and more frequently aggressive than male NAVs (Tarlow et al 2001:4). Subsequently, they are also more often the indirect cause of the chick’s death. To see examples of males acting extremely aggressively, we have to look more closely at mate rotation. In the Nazca boobies, mate rotation is the female’s choice. Sometimes females stay with their breeding partner, but often they will switch to a new one. This allows females to breed with recent non-breeding males that are in better physical and psychological condition. This also results in all females mating, while males go through seasons of breeding and non-breeding. The mate rotation takes one of two forms. Either, the female abandons the male and finds a new mate and a new nest site. Alternatively, the female finds a new mate and together the new mating pair force the old male out of the nesting site. In this instance, the male Nazca boobies do act aggressively as they are trying to protect their nest site. However, the female is on average 15% larger than the male and the male is outnumbered, so the new mating pair successfully takes control of the nest site (Maness and Anderson 2007: 6). Therefore, male NAVs are not more aggressive than female NAVs, and do not exhibit aggressive behavior towards chicks as a result of mate rotation, but rather exhibit this behavior towards their old mate and their new mating partner. Therefore, the hypothesis did not properly predict the reasons for this kind of NAV behavior, but there are other factors that lead to increased aggression. Figure 3: Graph showing the number of NAV events committed by failed breeders (Anderson et al 2004: 9) One reason that adult Nazca boobies exhibit NAV behavior is because of their own failed breeding attempts. As figure 3 shows, 88.2% of failed breeders engage in NAV behavior. (Anderson et al 2004: 10). Another reason for NAV behavior is victimization. Chicks that were more frequently victims of aggressive NAV behavior were significantly more likely to be aggressive NAVs themselves. During maltreatment, the chick experiences a five-fold increase in corticosterone levels. This cycle of the victim turning into the aggressor leads some people to refer to the phenomenon as a ‘cycle of violence’ (Grace et al 2010: 6). Figure 4: Graphs highlighting the difference between chicks that committed siblicide and ones that did not, and their subsequent frequency of committing NAV events and aggressive NAV events as adults (Müller et al 2007: 3) NAV behavior in Nazca Boobies has been compared to NAV behavior in their close relative the Blue-footed boobies. It has also been studied in relation to the practice of obligate siblicide. In Nazca boobies, 60% of fledglings commit siblicide. While committing siblicide, the chick’s testosterone levels sharply increase. Each graph in figure 4 shows two bars, one for 2ECA+ and one for 2EC-. 2ECA+ stands for nests where both eggs hatched and one sibling killed the other one. 2EC- are nests where only one egg hatched and siblicide was not committed by the surviving fledgling. As a result, in these nests the chick did not experience the sharp increase in testosterone levels. The graph shows that the chicks that committed siblicide are more frequent NAVs and more importantly, more frequently exhibit aggressive NAV behavior. This indicates a relation between committing siblicide as a fledgling, experiencing high testosterone levels during one’s youth, and aggressively attacking chick’s that are not one’s own as an adult (Müller et al 2007: 3). Figure 5: Androgen levels of Nazca and Blue-footed Booby hatchlings (Müller et al 2007: 3) Figure 5 shows androgen levels of Nazca (NZBO) and Blue-footed booby (BFBO) hatchlings. Blue-footed boobies do not engage in obligate siblicide and are born with lower androgen levels. Nazca boobies on the other hand hatch with higher androgen levels, show higher frequency of NAV behavior, and higher rates of aggressive NAV behavior (Müller et al 2007: 4). In conclusion, hypothesis 1 is not correct. Males are actually less aggressive than females. They exhibit aggression towards their old mate and her new mating partner when they are forced out of their nest, but there is no evidence to show that they then go on to be more aggressive towards chicks. Their aggression is aimed at defending their nest site, not taken out on other pairs’ chicks. There are three principle reasons for aggressive NAV behavior, that do not include the abandoned male’s frustration. Instead, the first reason is failed breeding attempts. Another reason is the Nazca booby’s own victimization as a chick, and the resulting exposure to high levels of corticosterone. Thirdly, participating in obligate siblicide and successfully killing one’s sibling, coupled with high androgen levels at birth and sharp increases in testosterone levels while engaging in siblicide, led to higher aggression levels in grown Nazca boobies. Hypothesis 2 To improve their own chances of finding a mate, NAVs injure young chicks to reduce the chick’s physical appeal once they are grown (reducing the attractiveness of the competition) Findings This hypothesis is somewhat correct. Since the Nazca boobies are long lived birds, the chick and the adult NAV will at one point in their lives compete for mates, and physically injuring them when they are young could reduce the chick’s fitness when it is grown.