Giant damselfish display more nest aggression towards Acapulco damselfish than conspecifics


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Giant damselfish display more nest aggression towards Acapulco damselfish than conspecifics

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Giant damselfish display more nest aggression towards Acapulco damselfish than conspecifics
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Las jaquetas gigantes en sus nidos son más agresivas contra las jaquetas Acapulco que contra individuos de su propia especie
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Abarca, Samantha V.
Ornburn, Camille A.
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Species of the family Pomacentridae, including damselfishes, provide parental care and show territorial behavior. Microspathodon dorsalis, Giant damselfishes, are known to be especially territorial within their nesting sites and will attack other species that go within a few meters of their nests. Another species known as Stegastes acapulcoensis, Acapulco damselfish, are often seen to inhabit near the Giant damselfish nests. From 11 May 2019 to 18 May 2019, we studied the aggressive behavior of Giant damselfish and Acapulco damselfish at multiple sites on the coast of Cuajiniquil, including Bajo Rojo, Bahía Thomas, and Jack’s Point. We observed how often Giant damselfish and Acapulco damselfish would attack each other and individuals within their own species. We also observed other factors that can influence aggressiveness such as location of nests, algae presence, the face color of Acapulco damselfish, and the number of damselfishes around each nesting site. Our results indicate that when near their nests, Giant damselfish are more aggressive than Acapulco damselfish. Furthermore, Giant damselfish chase away Acapulco damselfish more often than conspecifics. Also, Giant damselfish are more aggressive towards Acapulco damselfish with brown faces compared to Acapulco damselfish cream faces. This study gives insight to the aggressive behavior of two species of damselfish near Giant damselfish nests. ( , )
Abstract:
Especies de peces de la familia Pomacentridae, incluyendo las jaquetas, presentan cuidado parental y muestran comportamiento territorial. Las jaquetas gigantes (Microspathodon doralis), son conocidas por ser especialmente territoriales dentro de sus sitios de nidos y atacan otras especies que se acercan a pocos metros de sus nidos. Otra especie conocida como la Jaqueta de Acapulco (Stegastes acapulcoenis), se encuentran frecuentemente cerca de los nidos de las Jaquetas gigantes. Desde el 11 de mayo al 18 de mayo 2019 estudiamos el comportamiento agresivo de las Jaquetas gigantes y las Jaquetas Acapulco en tres sitios en la costa de Cuajiniquil: Bajo Rojo, Bahía Thomas y la punta de Jack. Nosotros observamos, con qué frecuencia las Jaquetas gigantes y las Jaquetas Acapulco se atacaban entre sí y con qué frecuencia atacaron individuos de su misma especie. Observamos otros factores que pueden influir en la agresividad además de la especie como la ubicación de los nidos, la presencia de jardines de algas, el color de las caras de las Jaquetas Acapulco, y el número de individuos por especie por sitio de nido. Nuestros resultados indican que cuando las Jaquetas Acapulco están cerca de los nidos de las Jaquetas gigantes, la Jaqueta gigante ataca las Jaquetas Acapulco más frecuentemente que las Jaquetas gigantes. Además, las Jaquetas gigantes persiguen las Jaquetas Acapulco afuera de su territorio, más frecuente que a sus conspecíficos. También, las Jaquetas gigantes son más agresivas que las Jaquetas Acapulco con las caras color café comparado a las caras color crema. Gracias a este estudio se entiende mejor la dinámica entre dos especies de Jaquetas cuando las Jaquetas gigantes están anidando.
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Aggressive behavior in two species of damselfish Abarca & Ornburn 1 Giant damselfish display more nest aggression towards Acapulco damselfish than conspecifics Samantha V. Abarca and Camille A. Ornburn University of California, Santa Cruz ; Department of Ecology and Evolutionary Biology University of California, Santa Barbara ; Department of Ecology, Evolution, and Marine Biology EAP Tropical Biology and Conservation Spring 2019 7 June 2019 ______________________________________________________________________________ ABSTRACT Species of the family Pomacentridae, including d amselfishes, provide parental care and show territorial behavior . Microspathodon dorsalis, Giant damselfishes, are known to be especially territorial within their nesting sites and will attack other species that go within a few meters of their n ests. Another species known as Stegastes acapulcoen s is, Acapulco damselfish, are often seen to inhabit near the Giant damselfish nests. From 11 May 2019 to 18 May 2019 , we studied the aggressive behavior of Giant damselfish and Acapulco damselfish at multi ple sites on the coast of Cuajiniquil, including Bajo Rojo, Bah ’ a Thomas, and Jack's Point . We observed how often Giant damselfish and Acapulco damselfish would attack each other and individuals within their own species. We also observed other factors that can influence aggressiveness such as location of nests, algae presence, the face color of Acapulco damselfish, and the number of damselfishes around each nesting site. Our results indicate that when near their nests, Giant damse lfish are more aggressive than Acapulco damselfish. Furthermore, Giant damselfish chase away Acapulco damselfish more often than conspecifics. Also, Giant damselfish are more aggressive towards Acapulco damselfish with brown faces compared to Acapulco dams elfish cream faces. This study gives insight to the aggressive behavior of two species of damselfish near Giant damselfish nests. ______________________________________________________________________________ La s Jaqueta s gigante s en sus nidos son m‡s agre sivas contra las Jaquetas Acapulco que contra individuos de su propia especie RESUMEN Especies de peces de la familia Pomacentridae, incluyendo las jaquetas, presentan cuidado parental y muestran comportamiento territorial. Las jaquetas gigantes ( Microspathodon doralis ), son conocidas por ser especialmente territoriales dentro de sus sitios de nidos y atacan otras especies que se acercan a pocos metros de sus nidos. Otra especie conocida como la jaqueta de Acapulco ( Stegastes acapulcoenis ), se encu entran frecuentemente cerca de los nidos de las Jaquetas gigantes. Desde el 11 de mayo al 18 de mayo, 2019 estudiamos el comportamiento agresivo de las Jaquetas gigantes y las Jaquetas Acapulco en tres sitios en la costa de Cuajiniquil: Bajo Rojo, Bah’a Th omas y la punta de Jack. Nosotros observamos, con quŽ frecuencia las Jaquetas gigantes y las Jaquetas Acapulco se atacaban entre s’ y con quŽ frecuencia atacaron individuos de su misma especie. Observamos otros factores que pueden influir en la agresividad adem‡s de la especie como la ubicaci—n de los nidos, la presencia de jardines de algas, el color de las caras de las Jaquetas Acapulco, y el nœmero de individuos por especie por sitio de nido. Nuestros resultados indican que cuando las Jaquetas Acapulco e st‡n cerca de los nidos de las

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Aggressive behavior in two species of damselfish Abarca & Ornburn 2 Jaquetas gigantes, la Jaqueta gigante ataca las Jaquetas Acapulco m‡s frecuentemente que las Jaquetas gigantes. Adem‡s, las Jaquetas gigantes perseguien las Jaquetas Acapulco afuera de su territorio, m‡s frecuente que a sus c onespec’ficos. TambiŽn, las Jaquetas gigantes son m‡s agresivas que las Jaquetas Acapulco con las caras color cafŽ comparado a las caras color crema. Gracias a este estudio se entiende mejor la din‡mica entre dos especies de jaquetas cuando las Jaquetas gi gantes est‡n anidando. ______________________________________________________________________________ INTRODUCTION Many organisms show parental care in different ways. This includes protecting their offspring through food provision, offspring brooding, offspring attendance, building nests, and more (Royle et al. , 201 3 ). There are multiple be nefits of parental care that will ultimately increase the fitness of the organism (Klug and Bonsall 2014). By providing parental care, the survival, growth and qualit y of offspring can increase (Klug and Bonsall 2014). Parental care has also evolved in close relation to other forms of social behavior, such as competition, mate attraction, mating, and group living (Royle et al., 201 3 ). This is evident among vast numbers of animals but is exemplified by comparisons among closely related fish. There are multiple forms of parental care in fish species , such as hiding their eggs, building their own nests, incubating their young internally, and/or guarding their nests (Balsh ine and Sloman 2011). The most predominant form of parental care in fishes is guarding , which often involves aggression towards egg predators, both conspecifics and heterospecifics (Gross and Sargent 1985). Aggression of these fish allow them to be more successful in protecting their nests from potential predators (Steinhart et al., 2004). This method of protecting offspring is seen in multiple species of damselfish, more specifically, Microspathodon dorsalis and Stegastes acapulcoen s is . Giant damselfish and Acapulco damselfish are in the same family, Pomacentridae . Giant damselfish are identified by their dark blue body and their lighter blue head (Robertson and Allen 1994). Nuptial males of this species have a very pale, almost white anterior half of the ir body and juveniles are blue with scattered neon blue spots (Robertson and Allen 1994). Acapulco damselfish are overall brown gold with a lighter head (Robertson and Allen 1994). Some Acapulco damselfish have a brown head, while others have a cream face. Some researchers believe that those with brown heads are likely adults, while those with cream faces are juveniles ( Palacios Salgado and Rojas Herrera 2012 ). Adults of these two species inhabit rocky substrates and coral reefs and are herbivores that feed on attached algae from their own algal gardens (Hernandez Velasco et al., 2016). Both species are known to be territorial and can be aggressive towards other fish, when guarding their territories and nests. They are especially aggressive during mating season when they select a rock to build their nests on, while they are laying their eggs, and when protecting their nests before their eggs hatch (Montgomery 1980). They tend to be most aggressive towards their own species and tow ards other algae eating fish es that encroach on their territory and nests. Most damselfish have similar breeding and reproductive strategies. First, a male will select where he and his mate will build their nest , either on a horizontal or vertical rock, and clean it to create space . This is followed by courtship , then the female will attach her eggs onto the rock surface and the male will fertilize the eggs (DeLoach and Humann 2007). Males will then guard and clean their nests until their eggs hatch (DeL oach and Humann 2007). Giant damselfish and Acapulco damselfish are often found to inhabit and place their nests in close

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Aggressive behavior in two species of damselfish Abarca & Ornburn 3 proximity to each other. Acapulco damselfish often try to feed near the territories of Giant damselfish, and for this reason interspec ific and intraspecific competition takes place. During preliminary observations, we even noticed a group of Acapulco damselfish feeding from what seemed to be an abandoned Giant damselfish nest. We also noticed that the male Giant damselfish that were guar ding their nests often chas ed away both species of damselfish that went near their nests. In this study, we address the main question: How is the aggressive behavior of Giant damselfish and Acapulco damselfish influence d by the presence of Giant damselfi sh? We predicted that Giant damselfish would be more aggressive than Acapulco damselfish around their nests. We will also attempt to answer: Which damselfish species are the nesting Giant damselfish more aggressive towards? We predicted that Giant damselfi sh would attacks Acapulco damselfish more than conspecifics while defending their own nests. We will also address the question: How does the location of the Giant damselfish nest affect the presence of other damselfish and the aggressiveness of Giant damselfish? We predicted that there would be more damselfish near horizontal rock nests, since the y seem to be more noticeable, and therefore the Giant damselfish will be more aggressive at these nests. Lastly, we attempt to answer: Which Acapulco damselfish are more aggressive, and which are the Giant damselfish more aggressive towards? We predicted t hat Acapulco damselfish with brown faces would be more aggressive , and therefore be attacked more often by Giant damselfish, than those with cream faces . In this paper, damselfish will refer to the two species studied, Giant damselfish and Acapulco damself ish. METHODS AND MATERIALS Th is study took place at multiple sites along the coa s t of Cuajiniquil, Guanacaste, Costa Rica, which were chosen based on preliminary observations and sites used in a previous EAP student project done by Marks (2006). These sites inc l uded Bajo Rojo, BahÂ’a Tomas West and Jack's Point at Isla David. We decided to separate Bajo Rojo into two sites, Bajo Rojo #1 and Bajo Rojo #2 because these locations were located on different sides of the rock reef . We observed the social and a ggressive behavior of Giant damselfish and Acapulco damselfish near Giant damselfish nests. We focused on observations around Giant damselfish nests regarding the behavior of Acapulco damselfish and Giant damselfish. While at each snorkeling site , we identified Giant d amselfish nests based on the grey colored eggs either on a horizontal or vertical rock. Once the nest was located, we measured out a 1.5 meter square around the nest using a tape measurer, and then placed four painted rocks to mark the ar ea . We then took note of whe ther the nest was on a horizontal or vertical rock, if there was algae near the nest and if damselfish were feeding on the algae . Then, for five minutes, we observe d and record ed the number of interspecific and intraspecific att acks between Giant d amselfish and Acapulco d amselfish. We defined an attack as physical contact or chasing of another damselfish. We also recorded the number of Acapulco damselfish and Giant damselfish that entered the set area during the five minutes. We took turns recording data so that we could get consistent data at each nest and not have repeated data. We also recorded footage of the Giant d amselfish defending their nests using a GoPro attached to a large rock , to later compare data . Lastly, ANOVA tests were run to analyze the statistical significance of our data.

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Aggressive behavior in two species of damselfish Abarca & Ornburn 4 RESULTS Only data collect e d at Bajo Rojo #1, Bajo Rojo #2 and BahÂ’a Tomas will be referred to for all graphs and considered for statistical analysis . Jack's Point was not considered beca use only two nests were found here, not giving us a sufficient amount of data to show statistical significance. However, data points from Jack's Point were included in the table and graphs to show all data collected. Giant damselfish were significantly mo re aggressive than Acapulco damselfish. Giant damselfish also attacked Acapulco damselfish, particularly those with brown faces, significantly more often than conspecifics. Acapulco damselfish equally attacked Giant damselfish and conspecifics. Also, Acapu lco damselfish with brown faces were not significantly more aggressive than those with cream faces. Nest location was not a factor that affected presence of damselfish nor the aggressiveness of Giant damselfish. Table 1. Total n umber of attacks by both s pecies of damselfish and number of each species present at each site. Location # Giant damselfish attacks on Acapulco damselfish # Giant damselfish attacks on Giant damselfish # Acapulco damselfish attacks on Giant damselfish # Acapulco damselfish attacks on Acapulco damselfish # of Giant damselfish # of Acapulco damsel f ish Bajo Rojo #1 50 9 6 21 39 60 Bajo Rojo #2 47 9 4 3 28 58 Bah Â’ a Tomas 47 3 1 48 12 69 Jack's Point 34 2 2 13 7 14 TOTAL 178 23 13 85 86 201

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Aggressive behavior in two species of damselfish Abarca & Ornburn 5 Figure 1. Comparison of total average attacks of damselfish . At Bajo Rojo #1 and Bajo Rojo #2 , Giant damselfish were significantly more aggressive than Acapulco damselfish ( F i g. 1 Bajo Rojo #1 , ANOVA, n=11, F=6.52, p=0.02 ; Bajo Rojo #2 , ANOVA, n=15, F=13.45, p=0 .001 ). At Bah’a Tomas, there was no significant difference between the average number of Giant damselfish and Acapulco damselfish attacks ( Fig. 1 ANOVA, n=8, F=0.0007, p=0.98). Figure 2. The average attacks by Acapulco damselfish to wards each species of damselfish. -1 ! 0 ! 1 ! 2 ! 3 ! 4 ! 5 ! 6 ! 7 ! 8 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #total attacks/#individuals present! Aggressor Species! Giant damselÞsh! Acapulco damselÞsh! -0.5 ! 0 ! 0.5 ! 1 ! 1.5 ! 2 ! 2.5 ! 3 ! 3.5 ! 4 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #attacks/#individuals present! Species Attacked! Giant damselÞsh ! Acapulco damselÞsh!

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Aggressive behavior in two species of damselfish Abarca & Ornburn 6 At all three sites, the difference between average number of attacks by Acapulco damselfish towards either species of damselfish was not statistically significant ( F i g. 2 Bajo Rojo # 1, ANOVA, n=11, F=1.83, p=0.19 ; Bajo Rojo #2 , ANOVA, n=15, F=0.05, p=0.82; Bah’a Tomas , ANOVA, n=8, F=1.86, p=0.19 ). Figure 3 . The average attacks by Giant damselfish to wards each species of damselfish. At all three sites, the average number of attacks by Giant damselfish on Acapulco damselfish, was significantly greater than the average number of attacks on Giant damselfish ( F i g. 3 Bajo Rojo # 1 , ANOVA, n=11, F=12.18, p=.0023 ; Bajo Rojo #2 , ANOVA, n=15, F=9.27, p=.00 5 ; Bah’a Thomas , ANOVA, n=8, F=11.87, p=0.0039). -0.5 ! 0 ! 0.5 ! 1 ! 1.5 ! 2 ! 2.5 ! 3 ! 3.5 ! 4 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #attacks/#individuals present! Species Attacked! Giant damselÞsh ! Acapulco damselÞsh!

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Aggressive behavior in two species of damselfish Abarca & Ornburn 7 Figure 4 . The average number of Giant damselfish attacks at each type of nest location. At Bajo Rojo #1, there were only nests on horizontal rocks, none on vertical rocks. At Bajo Rojo #2 and Bah’ a Tomas , there was no significant difference between the number Giant damselfish attacks toward other damselfish at vertical or horizontal rock nests ( F i g. 4 Bajo Rojo #2 , ANOVA, n=15, F=0.19, p=0.67 ; Bah’a Tomas , ANOVA, n=8, F=0.15, p=.71 ). Figure 5. T he average number of damselfish es present at each nest location type . 0 ! 5 ! 10 ! 15 ! 20 ! 25 ! 30 ! 35 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #Giant damselÞsh attacks/#nests! Nest Location! Horizontal Rock! Vertical Rock! 0 ! 5 ! 10 ! 15 ! 20 ! 25 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! Average # damselÞsh present! Nest Location! Horizontal Rock! Vertical Rock!

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Aggressive behavior in two species of damselfish Abarca & Ornburn 8 At Bajo Rojo #2 and Bah’a Tomas , the re was no significant difference between the amount of damselfish present at either vertical or horizontal rock nests ( F i g. 5 Bajo Rojo #2 , ANOVA, n=15, F=0.31, p=0.59 ; Bah’a Tomas , ANOVA, n=8, F=1.78, p=0.23 ). Figure 6 . The average number of attacks by Acapulco damselfish based on their face color. At Bajo Rojo #1, Acapulco damselfish with brown face s attacked a significantly greater number of times than those with a cream face s (Fig.6 ANOVA, n=11, F=4.34, p=0.05). There was no significant differen c e at Bajo Rojo #2 and Bah’a Tomas in Acapulco average attacks based on face color ( F i g. 6 Bajo Rojo #2, ANOVA, n=15, F=1.45, p=0.24 ; Bah’a Tomas, ANOVA, n=8, F=0.20, p=0.67 ). -0.5 ! 0 ! 0.5 ! 1 ! 1.5 ! 2 ! 2.5 ! 3 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #attacks/# present! Color of Acapulco damselÞsh face! Brown! Cream!

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Aggressive behavior in two species of damselfish Abarca & Ornburn 9 Figure 7 . The average number of attacks by Giant damselfish towards Acapulco damselfish based on the Acapulco damselfish face color. At all three sites, the average number of Giant damselfish attacks on brown faced Acapulco damselfish was significantly greater than on cream faced Acapulco damselfish ( F ig.7 Bajo Rojo #1, ANOVA, n=11, F =5.04, p=0.04 ; Bajo Rojo #2, ANOVA, n=15, F=6.19, p=0.02 ; Bah ’ a Thomas, ANOVA, n=8, F=3.74, p=0.07). DISCUSSION At two locat i ons, Bajo Rojo #1 and Bajo Rojo #2, Giant damselfish are significantly more aggressive towards Acapulco damselfish than conspecifics, which supported our original prediction (Fig. 1) This can be explained b ecause Giant damselfish are very territorial and are es pecially protective when guarding their nests. If Giant damselfish are threatened by other fishes, then they will chase them away. Giant damselfish were more aggressive because they were guarding their own nests, while Acapulco damselfish were trying to fe ed from and prey on the nest eggs. At all three locations , the number of Giant damselfish attacks on Acapulco damselfish was significantly greater than the number of attacks on Giant damselfish (Fig. 3 ). This supports our original prediction of Giant damselfish attacking more Acapulco damselfish than conspecifics . While protecting their nests, Giant damselfish could be very aggressive, and this seems to have been the case at all three of these sites. Ba sed on our observations, the Acapulco damselfish were feeding very close to the Giant damselfish nests, which threatened the Giant damselfish, causing them to attack. In our preliminary observations , we also noticed Acapulco damselfish feeding on Giant dam selfish eggs . This could explain why Giant damselfish are threatened by Acapulco damselfish and often chase them away. Previous studies have shown that territorial damselfish selectively keep out egg predators and food competitors, which we saw occurring o ften in nesting areas (Albrecht 1969 , Low 1 971 , Ebersole 1977 as cited in Montgomery 1980). Based on these findings, we can infer that Giant damselfish are aggressive -1 ! 0 ! 1 ! 2 ! 3 ! 4 ! 5 ! 6 ! Bajo Rojo #1! Bajo Rojo #2! Bahia Tomas! Jack's Point! #attacks/#present! Color of Acapulco damselÞsh face! Brown! Cream!

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Aggressive behavior in two species of damselfish Abarca & Ornburn 10 towards Acapulco damselfish to prevent them from preying on their eggs and feeding from a round their nests. A possible explanation for the fewer conspecific attacks by Giant damselfish could be that in most cases, many of the Giant damselfish nearby were guarding their own nests. Therefore, they might have been more focused on guarding their o wn nests rather than preying on other Giant damselfis h nest s . Overall, Giant damselfish are significantly more aggressive towards Acapulco damselfish with brown faces than those with cream faces (Fig. 7) . This finding supported our original predicti on. We noticed that Acapulco damselfish with brown faces seemed to be more aggressive than those with cream faces and therefore would be chased away by Giant damselfish more often. However, based on our findings, Acapulco damselfish with brown faces were n ot significantly more aggressive than those with cream faces (Fig. 6). Another explanation could be that Acapulco damselfish with brown faces are more likely adults and therefore Giant damselfish are more threatened by them (Palacios Salgado and Rojas Herr era 2012). We also found that Acapulco damselfish equally a t tack heterospecifics and conspecifics. We originally predicted that Acapulco damselfish would attack conspecifics more often than Giant damselfish, however this was not supported by statistical analysis (Fig. 2). Nest location was also found to not have an effect on Giant damselfish aggression and presence of damself ish (Fig. 4 and 5). We originally predicted that there would be more damselfish around horizontal nests and therefore, Giant damselfish at these nests would be more aggressive. However, there was no significant difference between horizontal and vertical ro ck nests with respect to damselfish presence and Giant damselfish aggression. Lastly, we predicted that Acapulco damselfish with brown faces would be more aggressive than those with cream faces. However, we found there was no significant correlation betwee n Acapulco damselfish face color and aggressiveness (Fig. 6). In conclusion, we found that when guarding their nests, Giant damselfish are more aggressive than Acapulco damselfish. We also found that Giant damselfish chase away Acapulco damselfish more of ten than conspecifics. Giant damselfish are also more aggressive towards Acapulco damselfish with brown faces compared to those with cream faces. Contrary to our prediction, Acapulco damselfish equally attacked both Giant damselfish and conspecifics. Acapu lco damselfish with brown faces were not significantly more aggressive than those with cream faces. Lastly, nest location did not significantly affect the presence of damselfish and the aggressiveness of Giant damselfish guarding these nests. Further areas of research could include why Giant damselfish attack some Acapulco damselfish near their nests, but not others, regardless of their face color. Behavioral research studies on damselfish are important to understand why it is that they behave the way they do. This study gives insight into the territoriality and aggressiveness of Giant damselfish and Acapulco damselfish. It is important to understand how organisms behave and interact with their environment to better understand the living world. ACKNOWLEDGE MENTS Thank you to Frank Joyce for helping us throughout the whole process and having extreme patience with all studen t s at Cuajiniquil . We are very grateful to the Lara family for their hospitality and generosity. Also, thank you to Minor Lara , Minor Jr . Lara , and Anibal Lara for taking us out to our snorkeling sites each day and for their knowledge; without them we could not do this project. We are also thankful for Emilia Triana and Federico Chinc h illa for

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Aggressive behavior in two species of damselfish Abarca & Ornburn 11 their advice and support. Thank you t o our Cua jiniquil peers that provided advice, help, and laughs. Lastly , thank you to our homestay families for opening their homes to u s in Cuajiniquil. LITERATURE CITED Balshine, S., and Sloman, KA. (2011) Parental Care in Fishes. In: Farrell A.P., (ed.); Encyclopedia of Fish Physiology: From Genome to Environmen t, volume 1, pp. 670 677. San Diego: Academic Press. Benham, C. 2005. Eff ects of Reef Substrate and Anthropogenic Impact on the Distribution of Seven Damsel f ish ( Pomacentridae) in the Bah ’ a Cuaj iniquil, Costa Rica. EAP University of Califo r nia, Instituto Monteverde Fall 2005. [Unpubli s hed] DeLoach, N. and Humann, P. (2007). "Damselfishes." Reef fish behavior . 1st ed. Jacksonville, FL: New W orld Publ, pp.180 207. Gross, Mart R., and R. Craig Sarg ent. "The Evolution of Male and Female Parental Care in Fishes." American Zoologist , vol. 25, no. 3, 1985, pp. 807 Ð 822., doi:10 . 1093/ icb/25.3.807. Hern‡ndez Velasco, Arturo, et al. "Occurren c e of Holacanthus Clarionensis (Pomacanthidae), Stegastes Leucorus, and Stegastes Acapulcoensis (Pomacentridae) at Magdalena Bay, B.C.S., Mexico." Marine Biodiversity Records , vol. 9, no. 1, 2016, doi:10.1186/s41200 016 0062 1. Klug, Hope , and Bonsall, Michael B. "What are the benefits of parental care? The im portance of parental effects on developmental rate." Ecology and Evolution , 12: 2330 2351, 2014. Marks, L.M. 2006. Comparing Coastal Reef Fish Species Diversity and Richness in B a h’a Cuajiniquil, Costa Rica. EAP University of Califo r nia, Instituto Monte verde Spring 2006 . [Unpublished] Montgomery, Linn W., et al. "The impact of non selective grazing by the Giant blue damself i sh, Microspathodon Dorsalis, on algal communities in the Gulf of California, Mexico." Bulletin of Marine Sciences , 30: 290 303, 1980 Montgomery, W. Linn. "Mixed Species Schools and the Significance of Vertical Territories of Damselfis h es." Copeia , vol. 1981, no. 2, 15 May 1981, pp. 477 Ð 481., doi:10.2307/1444245. Palacios Sal g ado, Deivis S., Rojas Herrera, Agustin A., "Partial xan thism in a specimen of Acapulco major, Stegastes acapulcoensis (Teleostei: Pomacentridae), from the Tropical Eastern Pacific." Pan American Journal of Aquatic Sciences, 2012, 7(3):175 177 Peterson, Christopher W., et al. "Male Mating Success and Female Choice in Permanently Territorial Damselfishes." Bulletin of Marine Science s, vol.5 7, no. 3, 1995, pp.690 704(15)

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Aggressive behavior in two species of damselfish Abarca & Ornburn 12 Robertson, David Ross, and Allen, Gerald R. (1994). "Damselfishes." Fishes of the Tropical Eastern Pacific , by David Ross Robertson and Gerald R. Allen, University of Hawaii Press, pp. 189 Ð 190. Royle, Nick J., et al. The Evolution of Parental Care . Oxford University Press, 2013. Steinhart, Geoffrey B., et al. "Increased Parental Care Cost for Nest Guarding Fish in a Lake with Hyperabundant Nest Predators." Behavioral Ecology , vol. 16, no. 2, 2004, pp. 427 Ð 434., doi:10 . 1093/ beheco/ari006. ) Sun, D., Blomberg, S.P., Cribb, T.H. e t al. Coral Reefs (2012) 31: 1065. https://doi org.proxy.library.ucsb.edu:9443/10.1007/s00338 012 0929 5

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Aggressive behavior in two species of damselfish Abarca & Ornburn 13 Appendix A Ð Satellite Images of Data Collection Sites Figure 1. Satellite images of the data collection sites. Marker one indicates w here Bajo Rojo #1 and Bajo Rojo #2 are . Marker two shows Jack's Point, and marker three shows Bah ’ a Tomas. Figure 2. Close up satellite ima g e of Bajo Rojo. Marker one indicates were Bajo Rojo #1 is and marker two shows Bajo Rojo #2. 1 2 3 1 2

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Aggressive behavior in two species of damselfish Abarca & Ornburn 14 Figure 3. Close satellite image of Jack's Point, near Isla David. Marker one shows where Jack's Point is. Figure 4. Close satellite image of Bah ’ a Tomas Ð West. The marker shows where data was collected. 1 1


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