Vocalizations and associated behaviors of the white-nosed coati, Nasua narica

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Vocalizations and associated behaviors of the white-nosed coati, Nasua narica

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Title:
Vocalizations and associated behaviors of the white-nosed coati, Nasua narica
Translated Title:
Vocalizaciones de pizotes (Nasua narica) y sus comportamientos asociados
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Giampaoli, Jeremy
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Text in English

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Subjects / Keywords:
Coatis--Behavior ( lcsh )
Pizotes—Comportamiento ( lcsh )
EAP Fall 2017
EAP Otoño 2017
Costa Rica--Puntarenas--Monteverde Zone
Costa Rica--Puntarenas--Zona de Monteverde
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Abstract:
Vocalizations emitted by mammals in a social context can be linked to behavioral states. These different states may exhibit specific acoustic characteristics, such as long call duration, harmonics, and wide frequency range. My objective in this study was to describe the different vocalizations of White-Nosed Coatis (Nasua narica) according to their behavioral context. I recorded vocalizations of ten individual males and a social group of coatis in Monteverde, Costa Rica. Based on the associated behaviors, I grouped the calls into three behavioral states: aggressive, relaxed, and alarmed. After analyzing the acoustic characteristics of the vocalizations using spectrograms, I further discriminated the calls into sub-states. I quantitatively and qualitatively described a variety of vocalizations (chirps, squawks, squeaks, trills, and grunts). Each behavioral sub-state exhibited a significantly different vocalization, demonstrating a clear relationship between vocalization characteristics and associated behavior. The quantitative acoustic traits I found align with Morton’s motivational-structure rules: that aggressive vocalizations exhibit a lower minimum frequency and wider frequency range than those of non-aggressive contexts. Additionally, inter-individual variation was prominent and could provide a method of individual identification within the species. ( , )
Abstract:
Las vocalizaciones que los mamíferos producen en un contexto social pueden estar relacionadas con diferentes contextos de comportamiento. Los llamados asociados a estos contextos pueden presentar características acústicas específicas, por ejemplo, larga duración, armonía, y amplios rangos de frecuencia. Mi objetivo en este estudio fue describir los diferentes llamados de pizotes (Nasua narica) según su contexto de comportamiento. Grabé las vocalizaciones de diez machos individuales y un grupo social de pizotes en Monteverde, Costa Rica. Categoricé las llamadas en tres estados de comportamiento: agresivo, relajado, y asustado. Después de analizar las características acústicas de las vocalizaciones utilizando espectrogramas, pude agrupar los llamados en sub-estados. Cada sub-estado presentó una vocalización considerablemente diferente, exponiendo una relación clara entre las características de las vocalizaciones y los comportamientos asociados. Las características acústicas cuantitativas que encontré siguen las reglas de estructura motivacional de Morton, las cuáles predicen que las vocalizaciones agresivas exponen una frecuencia mínima más baja y un rango de frecuencia más amplio que vocalizaciones en contextos no agresivos. Por otro lado, encontré una gran variación inter-individual, lo cual podría facilitar la identificación de individuos dentro de la especie.
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Student affiliation: Department of Mechanical and Aerospace Engineering, University of California, Los Angeles

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Monteverde Institute
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Monteverde Institute
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M39-00642 ( USFLDC DOI )
m39.642 ( USFLDC Handle )

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Giampaoli 1 !"#$%&'$(&")* !"#$ !""#$%&'() !"#$%&'()*'+*,#"*-#&," !"#$%& !"#$%&' !"#$"%&"'()" Jeremy Giampaoli Department of Mechanical and Aerospace Engineering University of California, Los Angeles EAP Tropical Biology and Conservation Program, Fall 2017 15 December 201 7 #$%&'()& Vocalizations emitted by mammals in a social context can be linked to behavioral states These different states may exhibit specific acoustic characteristics such a s long call duration harmonics, and wide frequency range. My objective in thi s study was to describe the different vocalizations of White Nosed Coatis (Nasua narica) according to their behavioral context. I recorded vocalizations of ten individual males and a social group of coatis in Monteverde, Costa Rica B ased on the associated behaviors, I grouped the calls into three behavioral states: aggressive, relaxed, and alarmed. After analyzing the acoustic characteristics of the vocalizations using spectrograms, I further discriminated the calls into sub states. I quantitatively and qualitatively described a variety of vocalizations (chirps, squawks, squeaks, trills, and grunts). Each behavioral sub state exhibited a significantly different vocalization, demonstrating a clear relationship between vocalization characteristics and asso ciated behavior The quantitative acoustic traits I found align with Morton's motivational structure rules: that aggressive vocalizations exhibit a lower minimum frequency and wider frequency range than those of non aggressive contexts. Additionally, inte r individual varia tion was p rominent and could provide a method of individual identification within the species !"#$%&'$#&"()*+,)+-&'".)*+/ !"#$"%&"'()" !"#"$%$" !"#$"%&'#()*&"+,'+"!('-"+ *+%,-+. Las vocalizaciones que los mam’feros producen en un contexto s ocial pueden estar relacionadas con diferentes contextos de comportamiento. Los llamados asociados a estos contextos pueden presentar caracter’sticas acœsticas espec’ficas, por ejemplo, larga duraci—n, armon’a, y amplios rangos de frecuencia. Mi objetivo en este estudio fue describir los diferentes llamados de pizotes ( Nasua narica ) segœn su contexto de comportamiento. GrabŽ las vocalizaciones de diez machos individuales y un grupo social de pizotes en Monteverde, Costa Rica. CategoricŽ las llamadas en tre s estados de comportamiento: agresivo, relajado, y asustado. DespuŽs de analizar las caracter’sticas acœsticas de las vocalizaciones utilizando espectrogramas, pude agrupar los llamados en sub estados. Cada sub estado present— una vocalizaci—n considerabl emente diferente, exponiendo una relaci—n clara entre las caracter’sticas de las vocalizaciones y los comportamientos asociados. Las caracter’sticas acœsticas cuantitativas que encontrŽ

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Coati Vocal izations and Associated Behaviors Giampaoli 2 siguen las reglas de estructura motivacional de Morton, las cu‡les pr edicen que las vocalizaciones agresivas exponen una frecuencia m’nima m‡s baja y un rango de frecuencia m‡s amplio que vocalizaciones en contextos no agresivos. Por otro lado, encontrŽ una gran variaci—n inter individual, lo cual podr’a facilitar la identi ficaci—n de individuos dentro de la especie. Mammals produce vocalizations in both intra and inter species communication M ore than four thous and species of mammals vocalize for a myriad of reasons, such as mating, territory protection, and warning signal s (Fitch 2006) These vocalizations can be analyzed with respect to behavior. A hypothesis presented by Eugene Morton attempted to link motivation, or behavior, with the acoustic structure of mammal and bird vocalizations (Morton 1977). Many studies follo wed, supporting the hypothesis that sounds emitted in aggressive contexts exhibit lower frequency than those in non aggressive contexts (August & Anderson 1987, Compton et al 2001). These motivational structure (MS) rules illustrate an acoustic difference between aggressive and non aggressive behaviors, yet there is large variance within the two behavioral contexts. The complex social interactions of white nosed coatis, Nasua narica necessitate a variety of acoustic communication (Gompper 1997). C oatis ma intain a social structure formed of female and juvenile groups, calle d bands, and individual males. Males leave the bands at two years of age to become solitary and territorial, rejoining a band only for a short mating period (Jenzen 1983). Both bands and individual males compete for food, foraging for approximately 90% of the day. (G ompper 1996, Wainwright 2007). The time spen t f oraging and competing for resources comprises much of the interaction between coatis, leading to communication in both aggressive and non aggressive contexts (Blevins 2000). Other researchers have investigated coati vocalizations and shown that these communications display different acoustic characteristics in different contexts according to MS rules. One study characterized two k inds of coati vocalizations: chirps (non aggressive) and squawks (aggressive). Chirps are short duration, high frequency, harmonic calls ; s quawks exhibit lower frequency, a wide r frequency range, and longer duration (Compton et al 2001) Anothe r study focu sed on the chirps describing the vocalization in depth and determining the purpose to be communication during movement (Maurello 2000) Outside of these two studies coati vocalizations have been described as barking, chirping, growling, grunting, his sing squealing, and chuckling. These labels are ambiguous without quantitative descriptions. Additionally, all previous coati vocalization studies have focused on a small group of captive individuals From observations and acoustic analysis I have found that N. narica individuals in the wild readily produce a wider variety of sounds and exhibit more behavioral states than previously studied individuals in captivity I n the present study I have created a more comprehensive catalogue of coati vocalizations, de scribed quantitatively and grouped by behavioral context The vocalizations exhibit inter individual variation as well as align with previous studies supporting MS rules in coatis.

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Coati Vocal izations and Associated Behaviors Giampaoli 3 /(&+'0(1%"2"/+&345% I studied ten solitary male individuals as well as a ba nd of females and juveniles between 20 November 2017 and 2 December 2017 on the grounds of the Estaci—n Biol—gica Monteverde (E BM ) in Puntarenas, Costa Rica At the EBM there is a population of coatis that are constantly attracted to the Station building because of the organic waste These coatis that inhabit in areas with high human impact are most active from 08:00 to 10:00 and 12:00 to 14:00 (Sabatke 2015 personal observation ). A fter breakfast, 08:00, and after lunch, 13 :00, I brought organic waste to a flat area on a trail to the southwest of the EBM to attract a group of coatis. 6789:6;<69="6976>67? #@A I had to identify each coati individually to ensure consistent data collection Be fore or during recording, I focused on the animals' face s and oth er traits to discover distinguishable characteristics Coatis often injure each other during aggressive encounters sometimes leaving distinguishing marks (Wainwright 2007 personal observations ). I discovered ear shape and imperfections, likely sustained from fights to be the most effect ive w ay of identifying individuals; overall coat color or patchiness was also important for differentiation I name d the individuals and document ed their distinguishing features with photographs on a Canon 1D Mk. IV DSLR Camera ( Appendix 1 ). B8C#>6D* 6#@"#9#@
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Coati Vocal izations and Associated Behaviors Giampaoli 4 Table 1: Behavioral states and sub states of coatis and distinguishing behavior of the individuals in each context. !"#"$ % &''()!!*+) % (),&-). % &,&(/). % 0$1#2345 % barring of teeth, upward pointed snout, biting, chasing sniffing, foraging, restin g, unaffected movement, lowered tail erect tail, cautious movement, retreating, alert eyes !67 8 !"#"$ % 9$553"453#:3;< % 915$#" % =4>?:3@" % A45#B3>B % *>C63;3"32$ % D4>$ % 0$1#2345 % vocalization towards a group, remaining within a distinct area vocalization towards a s pecific individual, without physical conflict physical conflict between two or more individuals feeding and searching for food friendly or expectant activity None *8ED*769=">DE#@6F#:6 D9A I record ed the vocalizations with a portable audio recorder, Olympus LS 12 Linear PCM Recorder, using an external Rode NTG2 Dual Powered Directional Condenser Microphone. During the two observation hours of each individual, I capture d the known chirps and squawks as well as previously unidentified vocalizations. Human in trodu ced food has been shown to increase coati aggressive interaction (Blevins 2000). Using the organic waste from the EBM I induce d contact between the male individuals to record noises made in an aggressive context. T wo total hours of observation spl it between multiple days and times of day, provide d enough time for me to record the coatis during their relaxed behavior. I elicit ed alarmed behavior by moving near an individual. #9#@DE#@6F#:6D9A I cut the raw audio files into clips that encompas sed one vocalization or one set of similar continuous vocalizations The behavioral observations recorded during the research allowed me to place each sound clip into a behavioral state and sub state (if applicable) as determined by t he distinguishing cha ra cteristics (T able 1). I then analyze d the audio record ings with Raven Pro 5 software. For each clip, I calculated the low frequency (Hz), high frequency (Hz), delta frequency (Hz), and delta time (s).

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Coati Vocal izations and Associated Behaviors Giampaoli 5 *+%,1&% Through the 20 hours of monitoring, I reco rded ten individual males as well as one female juvenile band. In total, I recorded 121 distinct, high quality recordings of coati vocalizations (T able 2) Table 2 : Total amount of audio clips per individual and per behavioral state. Band denotes t he soci al band of females and juveniles. *>E323E6#:% 45%'546F % &''()!!*+) % (),&-). % &,&(/). % *>E323E6#:% 94"#: % 9$553"453#: % 915$#" % =4>?:3@" % A45#B3>B % *>C63;3"32$ % % % &5"654 % # $ % $ % GH % 0#>E I % %& % JJ % =#?$@3"4 % ( & # GH % =#5#F3;"4:# % # ( %% ) % JK % =: 42$5 % & ) % K % *LLM % & & % % N % O$E#L4 % # " H % O$F$ % & " $ P % O$F$@3"4 % % % % ) H % O35#"# % ) ) % H % Q*/ % ) ) & P % 94"#: % R % SN % JP % JT % GT % G % GJG % I created a catalogue of coati vocalizati ons in each behavioral context. The stat es and sub state s exhibit significantly different vocalization characteristics Each state and sub state is described quantitatively (T able 3) and qualitatively. Table 3: Quantitative description of vocalizations for each behavioral context. !"#"$ % !67 8 !"# "$ % +4@#:3L#"34> % ,4U%A5$C% VWLX % W3B1%A5$C % V WLX % .$:"#%A5$C% VWLX % .$:"#% 93<$%V;X % !#39?5/@2 AB84CD %#&:;!! #(#';(! $)));(! !;)! )( E-54F-G H3,4+/9+ >=/,I %<%!;&! &'%);)& %
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Coati Vocal izations and Associated Behaviors Giampaoli 6 #==*8AA6>8" G :8**6:D*6#@6A/ H =*?9: I Territorial behavior is a form of aggressive behavior in which an individual exhibits space related dominance (Kaufmann 1983). In this study, threats and physical conflicts occupy their own respective sub states; the behavior specific to this sub state is directing vocalizations at multiple individuals and defensive behavior of a small region. The small region usually corresponded to the area with the organic waste. Only two male individuals consistently exhibited territorial behavior: Pepe and Carapistola. When behaving territorially, both males emitted grunts (Figu re 1 a ) Grunts are vocalizations with a wide frequency range with a r elatively small low frequency (T able 3) These vocalizations are short duration repeating at an average of 5 times per second with approximately 0.1 second time gaps between each grunt. Carapistola also emitted a territorial vocalization that can be recognized by human auditory discernment by vaguely resembling a dolphin call (F igure 1b ) The individual parabolic chirps are like those in the relaxed foraging behavioral state, but repeate d more frequently (every 0.1 seconds) and with increasing maximum frequency. Each set of vocalizations contains 8 to 15 repeated syllables. In my observation, no other individuals vocalized in a similar manne r. I recorded another unique vocalization i n thi s category from Pepecito that resembles a human scream (F igure 1c appendix 2 ) The individual was standing over a pile of organic waste and seemed to vocalize towards a group of other males within 3 meters.

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Coati Vocal izations and Associated Behaviors Giampaoli 7 Figure 1: a. Spectrogram of normal territori al aggressive grunts b. Spectrogram of Carapistola's territorial chirps followed by grunting. c. Spectrogram of unique territorial vo calization emitted by Pepecito #==*8AA6>8" G :C*8#: H:*6@@I I recorded more vocalizations in the threatening aggressive su b state than in any other behavioral cate gory (T able 2). This lead to a wide variety in threat vocalizations. Many of the sounds were characterized by quavering or vibrating between multiple frequencies. I classify t hese vocalizations as a trill (F igure 2 ) which has 2 or more peak s and multiple resonance bars. The duration of these sounds is relatively long and the frequency is relatively low (T able 3).

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Coati Vocal izations and Associated Behaviors Giampaoli 8 Figure 2 : a. Spectrogram of a trill from a member of the female juvenile band wi th two distinct pea ks. b. Spectrogram of a trill from Arturo with two distinct peaks. Within the female and juvenile band, there is not an extremely clear distinction between aggressive threatening and aggressive confl ict. Though the interaction (F igure 3) displays sim ilar audio characteristics to a squawk, this aggressive interaction did not result in physical conflict. For this behavioral characteristic, these vocalizations in the threatening aggressive state. An important distinction between these intra band threats and male versus male conflict is the length of the vocalization. Intra band threats lasted for an average of 1.32 seconds, as compared to the average 4.05 seconds of the male conflicts.

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Coati Vocal izations and Associated Behaviors Giampaoli 9 Figure 3 : Spectrogram of a threa t within a female juvenile band. #==*8 AA6>8" G ED9;@6E: HAJ?#KLI Aggressive behavior with physical contact often follow threats The vocalizations associated with physically aggressive interactions are termed squawk s (Compton et al 2001). Each squawk syllable is an inverted parabolic frequency curve (F igure 4) The syllable frequency shape can vary slightly based on individual and location within the call. Some squawk elements have two peaks but display t he same audio characteristics. The low frequency is slightly higher than the other two a ggressive sub states, but lower than the relaxed and alarmed contexts (T able 3). The length of a physically aggressive e ncounter varies with some interactions as short as 2.8 0 seconds and some as long as 10.3 3 seconds, averaging at about 4.05 second s. Figure 4 : Spectrogram of an aggressive encounter with physical contact. For clarity, only one coati is shown vocalizing *8@#M87" G ;D*#=69= Although the foraging behavior leads to many ag gressive in teractions, the act of foraging is a relaxe d, normal acti on for coatis (Gompper 1996). Coati chirps are short duration vocalizations with a small frequency rang e and four to seven resonance s (T able 3). The individual syllables are inverse parabolic frequency curves, as in squawks, yet are not repeated at a norma l i nterval (F igure 5). Some chirps are accompanied by audible sniffs.

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Coati Vocal izations and Associated Behaviors Giampaoli 10 Figure 5 : Spectrogram of normal foraging relaxed chirps *8@#M87" G 69J?6A6:6>8 Vocalizations emitted in the relaxed inquisitive context or squeaks, sound like chirps to the human ear yet display different auditory characteristics. Squeaks are much higher in primary frequency and al most twice as long on average (T able 3). The end of the frequency curve is much higher than the beginning, resulting in a squeak increasing in frequency. Figur e 6 : Spectrogram of normal inquisitive relaxed chirp s #@#*/87 I was only able to record one set of vocalizations from the alarmed behavioral state (F igure 7) The series of three vocalizations are characterized by a short chirp, decreasing in freq uency and without resonance bars (T able 3), followed by a nasal sound with large frequency range like the territorial grunts.

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Coati Vocal izations and Associated Behaviors Giampaoli 11 Figure 7 : Spectrogram of alarmed chi rps and nasal vocalizations Additionally, I observed a moment of alarmed and alert behavior caused by a large branch breaking in the forest about five meters away from the coatis. During the next 35 seconds, the individuals stopped feeding, did not vocalize, and looked toward the origin of the sound. All individuals had returne d to relaxed beh avior after two minutes 70%),%%04. #==*8AA6>8" B8C#>6D* Conservation of energy rules explain much of the aggressive behavior and vocalization of coatis. In the territorial aggressive sub state, the t wo male individuals that consistently exhibited territori al behavior ( Pepe and Carapisto la) appeared to be the largest and most dominant males in the area. This seems to align with the definition of territoriality dominant individuals attempt to control most of the resources (Kaufmann 1983). Human provided organ ic waste creates a very different feeding environment than naturally foraging for food ( Blevins 2000) This may have impacted my recording of the territorial vocalizations i n male coatis induced by food. The surplus of food resources provided by the organi c waste reduce s the need for territoriality. Thus, according to conservation of energy, only the largest and most dominant males are willing to exert the energy for territorial behavior. Threatening aggressive vocalizations also seem to be determined by en ergy output rules. The purpose of a warning call is to prevent a fight By producing a trill individual s avoid expending unnecessary energy in a fight and additional vocalizations. Males compete for resources and must budget the energy used on physical conflict and squawks. On the other hand, the band of females and juveniles forage cooperatively. Because of this communal behavior they do not need to expend the same amount of energy in resource competition as individual males (Gompper 1996). Though th e

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Coati Vocal izations and Associated Behaviors Giampaoli 12 members of the band compete on a small scale over specific food items they do not need to exhibit dominant territorial behavior over the area (Kaufmann 1983). This leads to shorter aggressive interactions and less physical confl ict as compared with the behavior of individual males. Aggressive vocalizations have much lower average minimum frequencies than those of non aggressive contexts (T abl e 3). These calls are also "harsh" sound, or sounds that exhibit a wider frequency b andwidth in a single syllable (F i gures 1, 2, 4). The function of aggressive communication, primarily to obtain resources, has selected for these acoustic char acteristics (Morton 1977). In non aggressive contexts (both relaxed and alarmed), the vocalizations are much hi gher in frequency and tonal, sounds with a distinct frequency at a given time visualized as thin lines on a spectrogram. These acoustic characteristics due to behavioral contexts support Morton's MS rules and show similar findings to previous researchers (August & Anderson 1987, Maurello 2000, Compton et al 2001). *8@#M87" B8C#>6D* The values and characteristics of coati chirps I found align with those of previous studies. Chirps are presumably used as contact calls, defined as a non aggressive form of contact behavior invo lving exchange of acoustic signals (Maurello 2000). M y observations that many of these chirps were emitted within the family band support this hypothesis Individual males may also use contact calls to communicate non aggressively Although they are solita ry and presumably territorial, males may still communicate non aggressively with other familiar males to convey location of resources or presence of foreign individuals. In communicating such information, individuals ultimately hope to increase fitness by eliminating unnecessary energy expenditure on aggressive behavior. In addition to intra specific relaxed communication, I observed a behavioral context that has not been identified in previous studies This relaxed sub state is characterized by an inquisit ive or appeasing posture, prima rily induced by human contact. At the EBM the coatis in the area have learned that organic waste originates from the kitchen and is dumped on a trail to the southwest ( Maricel Cruz pers. comm. Marvin Hidalgo pers. comm. ). A s the coatis recognized me through visual or olfactory cues, they expected organic waste to be offered. The coatis appeared to use inquisitive vocalizations squeaks, as an inquiry or appeasement in hope of food. I did not find similar squeaks in any intra species communication further supporting my explanation of this sub state as most often caused by human interaction. #@#*/87 B8C#>6D* During the entirety of my observation of coatis, I was only able to record a single audio clip of alarmed vocalizations. This could be due to a variety of reasons, but the strongest explanation is the familiarization with human presence. Coatis are more active in highly trafficked areas of Monteverde ( Sabatke 2015 ). These individuals have learned to view the E BM as a sou rce of food and no longer fear human presence as much as

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Coati Vocal izations and Associated Behaviors Giampaoli 13 individuals found in areas with less human traffic. As they grow accustomed to human disturbances, coatis will exhibit less alarmed behavior 69:8* 6976>67?#@">#*6#:6D9 Though this study did not ex plicitly test for inter individual variation or its purpose, I qualitatively noticed individual variation when analyzing the spectrograms and in field observations. Each coati individual seems to have their own voice: specific audio characteristics exclusi ve to one coati, regardless of behavioral state. This variation is seen across all vocalizat ions and behavioral states, yet can be subtle or distinct. Chirps display inter individual variation in the difference in frequency between the first and second ha rmonics (Maurello 2000). Grunts emitted by the two individuals vary slightly by duration and frequency range. Similar slight acoustic differences could be quantitatively tested for in squawks and squeaks as well. Trills exhibit the most varied category of vocalizations each individual emits a trill with a different duration and number of frequency peaks. These inter individual variations may be linked to identifying factors of an individual, su ch as size, age, or aggression. In a social group this individ ualization is useful in communicat ion with chirps, or contact calls For example, mothers can recognize their children and members of the group can identify a foreign individual. As seen with the large variation in trills e mitting a vocalization that conv eys identifying factors is important for individual males. Expressing this information in a threat would allow an aggressor to determine if a phys ical conflict is necessary, decreasing the potential energy output of both individuals Additionally, vocali zations may vary geographically by population ED9E@?A6D9 Coatis produce a wide array of vocalizations, many which have not been previously described. Each vocalization exhibited a clear association with a given behavioral state or sub state. In my analysis of the behavioral states and their respective calls, aggressive vocalizations have a significantly lower frequency than non aggressive calls. These quantitative acoustic characteristics support MS rules and align with previous research of coati vocalizati ons. Prominent inter individual variation may be used for individual identification within the species The ability to distinguish individuals acoustically augments the coatis' complex social structure and allows for conservation of ener gy in aggressive i nteractions. Because of this complex social structure, coatis may exhibit more vocalizations and behavioral states than I was able to observe over this two week study, such as courtship displays, specific maternal care, or inter specific competition I ho pe this vocalization catalogue will provide a foundation for future studies to explore inter individual variation, identify potential geographic acoustic characteristics, and further investigate the relationship between vocalizations and behavior.

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Coati Vocal izations and Associated Behaviors Giampaoli 14 #)N.4O1+ 5P+-+.&% I would like to thank AndrŽs Camacho and Emilia Triana for their support, suggestions, and expertise as advisors. Johel Chaves, Katherine Gonzalez, and Moncho Calder—n provided research guidance even when they were busy with other responsibilitie s. Thank you to Mackenzie Kui for his editing expertise. The Estaci—n Biol—gica Monteverde (specifically, owner Marvin Hidalgo) provided the space to research and, esse ntially, the coatis in question. By far the most important people to my investigation are the cooks at the EBM Maricel Cruz Le’ton Eliza Gonz‡lez, Laly Chavarria Arguedas, Luisa Vega Garc’a, and Sonia Araya Cozuca. These ladies provided the organic waste, critical to the experiment, as well as unending moral support and laughter.

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Coati Vocal izations and Associated Behaviors Giampaoli 15 @0&+ '(&,'+"E0&+5 August, Peter V. & John G. T. Anderson. "Mammal sounds and motivational structure rules: A test of the hypothesis." Journal of Mammalogy 68.1 (1987): 1 9. Blevins, Christina. "Aggressive behavior in white nosed coatis, Nasua narica : Does food cause fighting?" Monteverde Tropical Biology, UCEAP IMV, Fall 2000 (2000). Chausseil, Monika. "Visual Same L Different Learning, and Transfer of the Sameness Concept by Coatis (Nasua Storr, 1780)." Ethology 87.1 L 2 (1991): 28 36. Compton L.A. J. A. Clarke, J. Seidensticker, and D. R. Ingrisano. Acoustic Characteristics of White Nosed Coati Vocalizations: A Test of Motivation Structural Rules ." Journal of Mammalogy 82.4 (2001): 1054 1058. Fitch, W. T. "Production of vocalizations in mammals." Visual Communic ation 3.2006 (2006): 145. Gompper, Matthew E. "Sociality and asociality in white nosed coatis (Nasua narica): foraging costs and benefits." Behavioral ecology 7.3 (1996): 254 263. Gompper, Matthew E., John L. Gittleman, and Robert K. Wayne. "Genetic relate dness, coalitions and social behaviour of white nosed coatis, Nasua narica." Animal Behaviour 53.4 (1997): 781 797. Janzen, Daniel H. Costa Rican Natural History ( 1983) : 478 480. University of Chicago Press. Kaufmann, John H "On the definitions and functio ns of dominance and territoriality." Biological Reviews 58.1 (1983): 1 20. Maurello, M. A., J. A. Clarke, and R. S. Ackley. "Signature characteristics in contact calls of the white nosed coati." Journal of Mammalogy 81.2 (2000): 415 421. Morton, Eugene S. "On the occurrence and significance of motivation structural rules in some bird and mammal sounds." The American Naturalist 111.981 (1977): 855 869. Sabatke, Megan. "Effect of human activity on the behavior of Central American agouti, Dasyprocta punctata, and the white nosed coati, Nasua narica ." Tropical Ecology and Conservation, CIEE, Summer 2015 (2015): 105 115. Wainwright, Mark. The Mammals of Costa Rica (2007): 279 284.

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Coati Vocal izations and Associated Behaviors Giampaoli 16 #QQ+.50R Appendix 1: Identifying characteristics of the ten male individuals r ecorded *>E323E6#: % E$>"3?M3>B%=1#5#@"$53;"3@; % Pepe Divet in left ear Cafecito Juvenile and brown/golden head Pirata Three slices on right ear Arturo Boxy protrusion on right ear UIM Juvenile, very dark head, less glazing on fur, unmarred ears Pepec ito Divet in left ear and slice on right ear Clover Multiple slices on both ears Carapistola Large compared to other males, umarred ears, very glazed fur Izzy Slice in right ear Pedazo Piece out of right ear Appendix 2: Quantitative description of Pe pecito's vocalization (figure 1c). !"#"$ % !67 8 !"#"$ % +4@#:3L#"34> % ,4U%A5$C% VWLX % W3B1%A5$C % V WLX % .$:"#%A5$C% VWLX % .$:"#% 93<$%V;X % !#

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