Knight 1 Captive Jaguar Behav ior and Best Practice Management Kathleen E. Knight University of California, Los Angeles Institute of the Environment and Sustainability UCEAP Tropical Biology & Conservation Fall 2016 16 December 2016 Abstract As the number of jaguars in captivity grows, the need to better understand the activity level a nd behavioral effects of captiv e jaguars increases By understanding the specific activities and behaviors of jaguars in captivit y, it is possible to better provide for the overall health of these animal in rescue centers, zoos, and wildlife sanctuaries This first part of this study consists of a comprehensive activity budget report, which assess es the activity and inactivity levels i n a male jaguar an d female jaguar at the Centro Rescate de Las Pumas i n CaÂ–as, Guanacaste, Costa Rica. These activity budgets were constructed through an observational study consisting of daytime early morning, and evening observations and night time came ra trapping. The second part of this study explores the effectiveness of various fo rms of environmental enrichment, and makes recommendations to encourage incorporation of knowledge about individual jaguars into forms of environmental enrichment designed t o improve the quality of life of captive jaguars Activity of the two jaguars was found to differ between each individual, and a higher level of activity during the late night and early morning but was overall much lower than that of jaguars in the wild a s reported by Rabinowitz and Nottingham in 1985 Different stimuli in the environmental enrichment study produced different scores of effectiveness, with commercial perfumes intended for human use and logs scented with male jaguar scent scoring the highest. This study recommends that rescue centers understand individual jaguars through activity budget reports and individual behaviors, and take measures to provide environmental enrichment for captive jaguars in order to increase the ir levels of activity and overall wellbeing Compartamientos de Los Jaguares Cautivos y AdministraciÂ—n de PrÂ‡cticas Mejores Resumen Con el aumento en el nÂœmero de jaguares que viven en cautiverio se incrementa la necesidad de entender mejor sus compo rtamientos. Entendiendo las actividades especÂ’ficas y sus comportamientos es posible proveer mejores condiciones para ÂŽstos animales en centros de rescate, zoolÂ—gicos y santuarios. La primera parte de mi estudio consistiÂ— en observar los periodos de activi dad, asÂ’ como de inactividad de dos jaguares, uno macho y una hembra, en el Centro de Rescate Las Pumas, en CaÂ–as Guanacaste, Costa Rica. ObservÂŽ directamente ambos jaguares durante el dÂ’a y durante la noche dispuse cÂ‡maras trampa. En la segunda parte de m i estudio explorÂŽ formas de enriquecimiento del ambiente de cada jaguar, haciendo recomendaciones para alentar y conocer formas de enriquecimiento del ambiente para mejorar la vida de jaguares cautivos. Las actividades de los dos jaguares fueron diferentes con un nivel de actividad mayor durante la noche tarde y temprano por la maÂ–ana. La actividad de los jaguares en cautiverio fue menor que lo informado para jaguares silvestres. En cuanto al enriquecimiento de su ambiente, los diferentes estÂ’mulos produje ron marcadores de efectividades diferentes; los perfumes comerciales y los troncos con olores de jaguar macho recibieron marcadores mÂ‡s altos. Con mi estudio recomiendo que los centros de rescate entiendan a cada jaguar individualmente a travÂŽs de reportes de sus actividades y comportamientos especÂ’ficos, y provean enriquecimientos del ambiente a cada jaguar cautivo, aumentando asÂ’ los niveles de actividad y los niveles de salud de los animales.
Captive Jaguar Behavior and Best Practice Management Knight 2 Introduction The decline of jaguars throughout Central America has rapidly increased within the last century. Valued for their pelts, jaguars faced a severe population decline after massive hunting activity, especially throughout the 1960s for the US and European fur market until the passage of the US Endangered Species Act and the establishment of CITES (Sanderson, et al, 2002) (Robinson and Redford, 1991). In addition to hunting, jaguars face threats due to habitat loss and human jaguar interaction. As only 30% of jaguars' orig inal habitat remains in Costs Rica due to deforestation and habitat fragmentation this animal's estimated density range from 0.74 to 11.8 individuals per 100 km squared continues to be a challenging factor in its survival as these fragmented areas can onl y support small populations (Maffei, et al, 2011). Oftentimes, jaguars are perceived a pests to livestock and believed to pose potential dangers to human by many communities, but scientists and conservationists argue that these fears are not true (Palacios 2011). Yet each year, jaguars are killed unnecessarily, with females leaving young cubs behind; oftentimes, these young cubs are the jaguars that end up in zoos, rescue centers, and other wild animal facilities. Thus, the jaguars we see in captivity ar e oftentimes much different than their wild counter parts as they do not exhibit many of the same behaviors as wild jaguars (Law, 2016). This begs the question: what do these animals do in captivity and what if anything can we do to improve their live s in regards to environmental enrichment ? In order to understand the behavior and activity levels of captive jaguars, it is important to understand the behavior and activity levels of jaguars in the wild. According to a study by Rabinowitz and Nottingham i n 1985, wild jaguars in Belize exhibited predo minantly nocturnal behavior, with males exhibiting a peak in traveling during t he early evening, maintaining high levels of acti vity throughout the night, and exhibiting inactive behavior primarily during midda y. F emales exhibited similar patterns of activity and inactivity with a slight tendency towards higher le vels of activity 60% of the time during a 24 hour period as compa red to males at 57% of the time. Rabinowitz and Nottingham also discovered that bo th male and female jag uars exhibited territorial behaviors defined by both olfactory and visual cues in the form of urine, feces, and scraping; however, it should be noted that females tended to occupy ranges that overlapped with males presumably to facil itate mating opportunities While wild jaguars must hunt their prey, defend their territory, and reproduce, many captive jaguars do not have the opportunity to do any of these activities. Environmental enrichme nt may be a form of stimulation that can reduc e boredom, discourage anxious behavior (such as pacing), and provide a better quality of life for captive jaguars. Some enrichment studies have involved varying the diets of jaguars between raw meat and live prey, encouraging olfactory enrichment through t he introduction of urine from other species into areas of the jaguar's enclosure, or providing different toys such as cardboard boxes and paper machÂŽ prey animals; many of these introduced stimulants produced a strong reaction in captive jaguars (Law 2016 ) (Viscarra et al, 2006) The goal of my study is to determine the activity budget s of jaguars in captivity the types of behaviors captive jaguars display, and to determine the effectiveness of d ifferent enrichment techniques. By determining a comprehensive activity budget for each individual jaguar and assessing which forms of environmental enrichment are most effective, it is possible to make recommendations for best practice management of capti ve jaguars at Centro de Rescate, Las Pumas. Methods and Materials The project took place over the course of two weeks (15 November 201 6 through 26 November 2016) at Centro del Rescate de Las Pumas, in CaÂ–as, Guanacaste in Costa Rica and involved the two adult jaguars at Las Pumas: a male jaguar named Rafa, age approximately 17 years ; and a female jaguar named Curubanda, age approximately 5 years Both jaguars arrived at Las Pumas at an early age: Rafa, as a young cub, and Curubanda at approximately a yea r old. After Rafa's mother was killed by hunters, he was turned over to Las Pumas by the police. Curubanda
Captive Jaguar Behavior and Best Practice Management Knight 3 was found wandering through a backyard, weak with facial injuries from an attack of unknown entity. She is cross eyed and suffers vision loss due to surviving this attack. Rafa (left), and Curubanda (right) Diagrams of enclosures (not to scale) : Rafa (left), and Curubanda (right) This project consisted of t wo parts: a 32 hour observational study of both ja guars (16 hours of observation for each individual) and exploratory experimental e nrichment studies with both jaguars. The dates for days of specific types of data c ollection are listed in the following table. Table 1. Dates and types of data collection
Captive Jaguar Behavior and Best Practice Management Knight 4 I. Activity Budget Study 1.Observational Study The first part of this activity budget study consisted of observations during the daytime, early morning, and the late afternoon/evening. Daytime o bservations of each jaguar were completed during the hours of 0800 and 1600 hours, with four hour long observation periods: two separate hour long periods in the morning, and two separate hour long periods in the afternoon. Daytime observations alternated between full days with each individual jaguar. I also complet ed two early morning observation days between the hours of 0500 and 0800 hours of both Rafa and Curubanda in addition to one night observation of Curubanda from 1600 to 2100 hours. T he scheduled night observation on 25 November 2016 for Rafa was canceled due to the effects of Hurricane Otto. I observed Rafa for an extra four hour period during the day, and thus the hour long observation on 15 November 2016 from 0800 to 0900 hours is include d in his early morning activity count. I observed from the outside of the enclosures, either in front of the visitor paths or from the forested area surrounding the enclosures D uring my observations, I noted each time the jaguar changed from its previous active or inactive state and recorded the time at which the change occurred. W alking or climbing, grooming, and eating defined active behavior; resting or sleeping defined inactive behavior The se different types of are discussed in the table below. Table 2. Types and description of activitie s categorized in observational study 2. Territorial and Aggressive Behavior In addition to observing and recording the changes in activities in order to complete an activity budget for each jaguar I also recorded the number of territorial or aggressive behavior displays during each hour observation period. I considered this in my study of captive jaguars in order to determine differences in ind ividual behavior and if either Rafa or Curubanda exhibited more or less displays when compared to one the othe r. The totals for both territorial and aggressive displays for all hours of daytime observat ion are represented by day number (numbers 1 3) and the totals for both types of behaviors for all hours of early morning or evening observations are represented by day number as well (numbers 4 5) Utilizing the standardized ethogram for big cats by Stanton et al in 2015, I categorized and quantified territorial and aggressive behaviors. Territorial displays were defined by : clawing, body ruba, (rubbing length of body against a modifier), head ruba (rubbing head against a modifier), urine spray (demarcation urination) and scratching trees inside the enclosure. Aggressive displays were defined by: baring teeth, striking at fence towards target charge, rushing the e nclosure fence toward a target, snarling, hissing, and growling 3 Camera Traps In order to account for nighttime activities from 16:00 hours to 08:00 hours, I utilized five Bushnell Trophy Camera HD camera traps which were placed on the fences surroun ding all sides of the enclosures. I alternated nights of camera trapping for each jaguar. I completed camera trapping of Rafa on the nights of 19 20 November 2016 and 23 24 November 2016, and completed camera trapping of Curubanda on the nights of 18 19 No vember 2016 and 20 21 November 2016.
Captive Jaguar Behavior and Best Practice Management Knight 5 In order to represent the activities of Rafa and Curubanda during the night, I counted the number of times the jaguar appeared on t he five camera trap s each hour. II. Enrichment Studies I performed enrichment studies to determine Rafa and Curubanda's reactions to different environmental changes and stimuli, with the intent of assessing how well captive jaguars responded to different changes. E very stimulant received a score of effectivene ss for each jaguar which was based off of the amount of the jaguar spent with each particular enrichment item I noted specific types of behaviors, including territorialism (defined in the "Territorial and Aggressive Behavior" section) over the item, purr ing, rubbing, and sniffing, but these were not included in the scoring system. I completed t wo differ ent types of enrichment studies with differe nt stimulants, described below, which all received scores based off of my scoring system, also described below. Table 3. Types of senses stimulated and corresponding abbreviations Table 4. Time spent with particular enrichment stimuli and corresponding s core of effectiveness 1. Perfumed Canvas Painting The first enrichment study was an observational study on the reactions of the jaguars to canvases prayed with Chanel No. 5 perfu me and then covered in paint. As this study combined three different types of sense stimulation (olfactory, visual and tactile), it was the most complex s timuli This was done by Las Pumas worker Ester, with the intention of selling the jaguar "art" at an upcoming Gala to raise money for the center. My observations noted the time spent with the perfumed canvases, and the specific types of behavioral reactio ns of each individual jaguar. 2. Environmental Introductions The second stimulant study was an observational and manipulative study u sing "toys" and scent marked logs from enclosures of other animals. With each stimulant, I recorded the time it took unt il the stimulant was discovered and how long the interaction with the stimulant lasted, in addition to observational notes on specific types of behaviors. Rafa received a toy made out of a hollowed coconut filled with straw from the rabbit and rat enclosures, and macaw and parrot feathers stuck into the sides. This was then strung inside his enclosure with a rope that could lower or raise the coconut toy. In addition, three logs from the ocelot enclosure and one log from the parrot enclosure were placed in various locations inside his enclosure. One ocelot log was placed in the back right corner of the enclosure where he spends time pacing. The other two ocel ot logs were placed next to the entrance of his food shelter and on top of his sleeping perch. The parrot log was placed on top of the sleeping perch and stretched across to the adjacent tree. In addition, Calvin Klein Obsession for Men perfume was placed at jaguar standing height on the tree in the grassy area.
Captive Jaguar Behavior and Best Practice Management Knight 6 Figure 2: Location of toy (A), logs with ocelot scent (B), log with parrot and macaw scent (C), and Obsession for Men by Calvin Klein perfume (D). Curubanda received two logs from the ocelot enclosure, and three logs taken from Rafa's enclosure. All of these logs were placed near the entrance to her food shelter and near the entrance to her enclosure (as it was not possible to fully enter the enclosu re). Figure 3: Location of logs with ocelot scent (B) and logs with male jaguar scent (E). Results I. Activity Budget Study 1. Observational Study During the daytime observational periods, Rafa exhibited walking or climbing active behavior every day, whereas Curubanda exhibited walking or climbing activity on only two of the days, and at considerably lower percentages. Whereas Rafa exhibited active walking or climbing behavior at a maximum of 37.6% of the time and a minimum of 19.6%, Curubanda exhibite d a maximum of 5.9% and a minimum of 0.42%. While Curubanda dedicated more time to inactive behavior (defined by resting or sleeping), the majority of her time was spent resting rather than sleeping on Days 1 and 2. Curubanda spent 96% of the day inactive, and only 4% of the day active; Rafa spent 69% of the day inactive, and 31% of the day active. The breakdowns of percentage of time for each individual active or inactive behavior are described in Figure 4(a). During the early morning and evening observa tional periods, Rafa's early morning activity budget averaged for Day 1 and Day 2 totaled 13.2% of the time active during the early morning, and 86.8% of the time inactive; Curubanda 's early morning activity budget totaled 33.5% of the time active and 66.5 % of the time inactive, which is the largest recorded total of active behavior
Captive Jaguar Behavior and Best Practice Management Knight 7 during any daytime, early morning, or evening period of observation between both Rafa and Curubanda. During the evening, Curubanda spent 17.9% of the time active and 82.1% of th e time inactive. The percentage of time for each different active and inactive behav iors are detailed in Figure 5. Figure 4: Percentage of time of each type of daytime activity and inactivity com pared between both jaguars. "Day" is defined by chronological completion of the observation period for each jaguar (the first day of observation described "Day 1," the second day described as "Day 2," and so on). "#$! %#&! '#&%! $! %#(! %#"! '#)"! "'#(! *+#(! $#+! %,#(! '#)"! ""#+! %$! "*#%! $+#(! %(#*! )*#%! $%#*! ()#"! $*#(! %*#%! "%! $,#+! '-! *'-! "'-! %'-! )'-! $'-! ('-! ,'-! &'-! +'-! *''-! *,./01.*(! *$./01.*(! *+./01.*(! *(./01.*(! "%./01.*(! "'./01.*(! 2343! 5676839:3! 2343! 5676839:3! 2343! 5676839:3! ;3! ?700@! A3BCDEBF@8! 7=G>! GB==H! "#$! "#$!
Captive Jaguar Behavior and Best Practice Management Knight 8 Figure 5. Percentage of time of each type of early morning or even ing activity or inactivity compared between both jaguars. 15 November 16 is comprised of one hour of morning observation, 18 November 16 is comprised of four hours of evening observation, and 21 and 22 November 2016 are comprised of three hours of morning observation s 2. Territorial and Aggressive Behavior Rafa exhibited a considerable higher number of territorial displays than did Curubanda, while Curubanda exhibited a significantly higher number of aggressive displays. On Day 3, Rafa exhibited the highest number of territorial displays (11 displays); the highest number of territorial displays exhibited by Curubanda was one (urine spray/demarcation urination) during the early morning observation period on 22 No vember 2016. Curubanda displayed the most aggressive displays out of the two jaguars, with 22 displays occurring on Day 3. Rafa's aggressive displays were significantly less, with the highest number of aggressive displays totaled at four for Day 2. (#%! (! %! "! *,#&! "$#"! )"#+! %"#)! %"#"! )*#,! $,#*! )+#,! )*#)! ")#&! +#(! '-! *'-! "'-! %'-! )'-! $'-! ('-! ,'-! &'-! +'-! *''-! *$./01.*(! *&./01.*(! "*./01.*(! ""./01.*(! 2343! 5676839:3! 2343! 5676839:3! ;3! GB==H! =3>! (#%! (! %!
Captive Jaguar Behavior and Best Practice Management Knight 9 Fig ure 6. Territorial and aggressive displays by Rafa for Days 1 3 (daytime observations) and Day 4 (morning observation 15 November 2016) and Day 5 (morning observation 21 November 2016). Figure 7. Territorial and aggressive displays by Curubanda for Days 1 3 (daytime observations) and Day 4 (evening observation 18 November 2016) and Day 5 (morning observation 22 November 2016). 3 Camera Traps Curubanda exhibited a high level of nocturnal activity, with five hour long periods representative of over 50 times captu red. There were only seven hour long periods in which Curubanda was not caught on camera, thus resulting in inactivity occurring only 21.9% of the 32 hours recorded, and activity occurring during 78.1% of the 3 2 hours recorded. Rafa did not exhibit a particularly high level of nocturnal activity in terms of number of times he appeared on the camera traps and did not have any hours in which capture number was in the range of 31 50 times or over 50 times Of the 32 hours recorded, Rafa was inactive for 12 hour long periods, thus resulting in inactivity occurring 37.5% of the time, and activity occurring 62.5% of the time. Table 5. Camera trap occurrences of each jaguar during the hours between 16:00 and 08:00. Ea ch number is representative of the number of times the jaguar appeared on the five camera traps. The hours between 18:00 and 06:00 are considered nighttime hours. '! $! *'! *$! "'! "$! '! *! "! %! )! $! (! ;6.7"#)*+)/-3<='03) /'0) 1'+'),"##-&*#-'=)'%8)9((#"33-2")/-3<='03) 2343!I=77F>07F3B!! 2343!J?7=GGF09! '! $! *'! *$! "'! "$! '! *! "! %! )! $! (! ;6.7"#)*+)/-3<='03) /'0) 56#67'%8'),"##-&*#-'=)'%8)9(#"33-2") /-3<='03) 5676839:3!I=77F>07F3B!! 5676839:3!J?7=GGF09!
Captive Jaguar Behavior and Best Practice Management Knight 10 4. Total Activity Levels The following chart provides a comprehensive average activity level for each jaguar during a 24 hour period. Rafa was active a total of 27.2 % during an averaged 24 ho ur period, and inactive for 72.8 %. Curubanda was active 33.4% during an averaged 24 hour period and inactive 66.6% Time Period Rafa Activity Percentage Curubanda Activity Percentage Daytime 31% 4% Early Morning 13.2% 33.5% Evening 17.9% Nighttime 37.5% 78.1% 24 hour total 27.2% 33.4% II. Enrichment Study As it was not possible to enter fully into Curubanda's enclosure, the ability to give her certain enrichment stimulants was not possible, and thus a full comparison between Rafa and Curubanda cannot be made. However, stimulant scores are reflective of indi viduals and thus the effectiveness of a particular stimulant varies with each jaguar. The most effective stimulants for Rafa were Obsession for Men by Calvin Klein and Chanel No. 5 and acrylic paint on canvas as they both received scores of 4. The most ef fective stimulants for Curubanda were Chanel No. 5 and acrylic paint on canvas and the logs with male jaguar scent, which received scores of 5 and 4, respectively.
Captive Jaguar Behavior and Best Practice Management Knight 11 Table 4 (a) Stimulant effectiveness for Rafa Table 4 (b) Stimulant effectiveness for Curubanda Discussion In this study I attempted to determine a comprehensive activity budget of captive jaguars and investigate the most effective enrichment techniques, with the goal of these observations being to develop best practice management stra tegies for captive jaguars. Several aspects must be included in a holistic approach to best practice management individual activity budgets, behavioral differences between jaguars, and the effectiveness of a variety of enrichment strategies. Activity bud gets provide one of the most comprehensive views of individualism in jaguars, in addition to providing valuable information to rescue centers regarding the active and inactive behaviors of their jaguars. Understanding the differences in diurnal and nocturn al activity preferences for individual jaguars may give rescue centers a basis by which to compare changes in activity level, and therefore lend further insight into the health of individual jaguars. While activity levels may vary between individual capt ive jaguars, I was able to determine activity budgets for both Rafa and Curubanda during the daytime, early morning, and evening
Captive Jaguar Behavior and Best Practice Management Knight 12 through an observational study, and during the night through camera trap capture analysis When considering all active and inactive percentages of time for each time period (daytime, early morning, evening, and nighttime), Rafa was active 27.2% of the time and inactive 72.8% of the time; Curubanda was active 33.4% of the time and inactive 66.6% of the time Comparing these results to Rabinowitz and Nottingham (1985), both Rafa and Curubanda were significantly less active than wild male and wild female jaguar activity averages. How ever, Curubanda was more active than Rafa, which is consistent wit h Rabinowitz and Nottingham's findings of higher activity levels in females than males, although this may be due to discrepancies in age rather than sex. Rafa exhibited the highest amount of active walking or climbing behavior between the hours of 12:00 an d 15:00, often in the form of pacing, presumably as a result of anticipated feeding. As wild males were active 57% of the time, and wild females were active 60% of the time, Rafa and Curubanda were 29.8% and 26.6% respectively less active than wild jaguars These differences may be explained by the lack of a variety of active behaviors not engaged in by most captive jaguars, for example hunting and mat ing, or by individual jaguar differences, such as Rafa's age or Curubanda's vision loss. Keeping in mind the difference s between Rafa and Curubanda's activity levels when compared to wild male and female jaguars, various environmental enrichment activities may increase the activity level of captive jaguars and reduce the difference in activity levels between captive jaguars and wild jaguars. By determining the effectiveness of different types of stimuli, it may be possible to increase the activity levels of jaguars in captivity. While this study did not incorporate a comparative difference between activity lev els during days of environmental introductions and days without environmental introductions, I did determine which environmental introductions and stimuli were most effective at encouraging active behavior based on time intervals. In both Rafa and Curuband a, Chanel No. 5 perfume received high scores of effectiveness, in addition to Obsession for Men by Calvin Klein introduced to Rafa, and logs with male jaguar scent introduced to Curubanda. Behavioral reactions of captive jaguars to both Chanel No. 5 and Ob session for Men by Calvin Klein were studied by Viscarra et al in 2006 and were both found to produce purring, rubbing, and sniffing reactions. While Viscarra et al focused on patterns of behavioral reactions in jaguars to determine effectiveness of the d ifferent perfumes, both this study and my own study indicated strong reactions to Chanel No. by captive jaguars. While the survey done by Law in 2016 did not include commercial perfumes for humans, it did include logs from other animals and toys from other cats as olfactory enrichment technique s, which received high scores based off of jaguar behavioral reactions; toys from other cats scored the highest possible score in Law's survey, while logs from other animals scored mid to high range While neither Rafa nor Curubanda produced significantly high scores in regards to logs from the ocelot enclosure or logs from the parrot and macaw enclosure, the introduction of logs with male jaguar scent produced a high score in Curubanda. Individual b ehavior is an important aspect in understanding captive jaguars. Territorial and aggressive displays by Rafa and Curubanda differed in their frequency and number, with Rafa exhibiting a higher number of territorial displays and fewer aggressive displays du ring the five days of data collection, and Curubanda exhibiting a higher number of aggressive displays, and only one territorial display in the five days of data collection. Rabinowitz and Nottingham (1985) found males to display more signs of territoriali sm, and this pattern appeared when comparing between Rafa and Curubanda. Understanding behavioral differences between individual captive jaguars can help a rescue center better provide for the needs of individuals; for example, higher signs o f aggression o r territorialism may provide insight into specific behaviors and personality differences between individuals. Having a comprehensive understand ing of individual jaguars increases the ability of a rescue center to provide for its animals. Another key compon ent to individual behavior includes understanding individual activity budgets. For example, while Rafa's daytime activity budget includes eating (1% of the day) as an active behavior, Curubanda's daytime activity budget does not. However, Curubanda's eveni ng activity budget includes eating as an active behavior (9.6% of the evening), and she was also observed eating at 22:30 hours on the camera traps. As both
Captive Jaguar Behavior and Best Practice Management Knight 13 jaguars were fed at the same time every day between 14:00 and 15:00 hours and within 15 minutes of each other, this provides further i nsight into individual behavior, indicating differences in consumption patterns between captive individual s Based on preliminary observations involving food searches done with Rafa, I would suggest further research int o the effectiveness of variegated food enrichment techniques. Hiding food throughout enclosures may encourage behaviors associated with hunting (utilization of olfaction, prey stalking moving and/or hiding food ) and increase activity of captive jaguars. I n addition, staggered feeding times may also serve to limit restless behaviors (pacing, growling) associated with scheduled meal times. As feeding is an integral part of a captive jaguar's day, there exist endless possibilities for future enrichment studie s in order to encourage hea lthy challenges and increased complexities for individuals. In addition to the enrichment stimuli that I provided for Rafa and Curubanda, I would recommend further investigation into effective enrichment strategies, keeping in mi nd the amount of time engaged in a stimuli and the positive behavioral reactions (purring, sniffing, rubbing, and territorialism). I recommend introduction of logs with female scent into Rafa's enclosure, bi weekly rotating introductions of Chanel No. 5, O bession for Men by Calvin Klein, and Margaret Astor Jovan Musk (as used in Viscarra et al, 2006) on different environmental introductions (such as cardboard boxes or hay) and on preexisting enclosure structures (such as logs on the ground or elevated in tr ees) in order to increase positive active behaviors. Environmental introductions and stimuli should be considered a key component to the maintenance of captive jaguars, as they increase the complexity of jaguars' days, encourage active beha vior, and contri bute to improving their quality of life by encouraging natural behaviors. Acknowledgements Many thanks to Martha, Rosita, Ester, Simon, and the rest of the staff at Las Pumas this project would indubitably not have been possible without your flexibility and willingness to allow me to turn into a crazy cat lady and to Julia the dog for snuggling on my lap when I checked camera traps in the mornings Thank you to my homestay family for accommodating my hectic schedule at Las Pumas, and for ensu ring my safety during early morning and late night observation times. Thank you to Federico for indulging in my jaguar obsession and providing me with so many wonderful resources and research papers. A huge thank you goes to Frank Joyce for enabling me to complete a project in a previously unexplored area, and for never failing to address the needs (and wild requests) of all of his students with an unwaveringly devoted attention. Lastly, thank you to the students and staff of EAP Fall 2016 you have all made this program an unforgettable experience Each and every one of you were an absolute joy to get to know and work alongside, and I am so proud of all of the things that we have accomplished.
Captive Jaguar Behavior and Best Practice Management Knight 14 Works Cited Law, C. "Environmental Enrichment." Jaguar Species S urvival Plan: Guidelines for Captive Management of Jaguars. American Zoo and Aquarium Association. 2016. 48 54. Maffei, L., Polisar, J., Garcia, R., Moreira, J., and Noss, A. J. "Perspectives from ten years of jaguar ( Panthera onca) camera trapping in Mes oamerica." Mesoamerica. Volume 15, no.1. August 2011. 49 59. Palacios, G., Cruz, E., and Guiris, M. "Current status of jaguars in Chiapas." Jaguar COnservation and Management in Mexico: Case Studies and Perspectives. 2011. 83 92. Rabinowitz, A. R,. and Nottingham, B. G. "Ecology and behaviour of the Jaguar ( Panthera onca) in Belize, Central America." Journal of Zoology. 149 159. Robinson, John G., and Kent H. Redford. Neotropical Wildlife Use and Conservation. The University of Chicago, 1991. Sanders on, E. W., Redford, K. H., Chetkiewicz, C. B., Medellin, R. A., Rabinowitz, A. R., Robinson J. G., and Taber, A. B. "Planning to Save a Species: the Jaguar as a Model." Conservation Biology. Volume 16, no. 1. Febr uary 2002. 58 72. Stanton, L. A., Sullivan M. S., and Fazio, J. M. "A standardized ethogram for the felidae: A tool for behavioral researchers." Applied Animal Behaviour Science. Volume 173. December 2015. 3 16. Viscarra, M. E., Ayala, G., Wallace, R., and Nallar R. "The use of commercial perfumes for jagaurs." Cat News. Volume 54. Spring 2011.