The Human-Alloprimate Interface in Gandoca, Costa Rica: an Ethnoprimatological Approach to Assess Conflict Between Residents and Three Neotropical Primates

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2017 The umh an-alloprimate interface in Gandoca, Costa Rica: an ethnoprimatological approach to assess conflict between residents and three Neotropical primates Giselle M. Narvaez Rivera Iowa State University

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The human-alloprimate interface in Gandoca, Costa Rica: an ethnoprimatological approach to assess conflict between residents and three Neotropical primates

Giselle M. Narváez Rivera

A thesis submitted to the graduate faculty

in partial fulfillment of the requirements for the degree of

MASTER OF ARTS

Major: Anthropology

Program of Study Committee: Jill D. Pruetz, Major Professor Cassandra Nuñez Maximilian Viatori

Iowa State University

Ames, Iowa

2017

TABLE OF CONTENTS

Page

LIST OF FIGURES ...... iv

LIST OF TABLES ...... v

NOMENCLATURE ...... vi

ACKNOWLEDGMENTS ...... vii

ABSTRACT………………………………...... viii

CHAPTER I INTRODUCTION AND LITERATURE REVIEW ...... 1

Ethnoprimatology ...... 1 Crop-raiding by nonhuman primates ...... 3 Why study human attitudes and perceptions towards nonhuman primates? ...... 6 Nonhuman primates of Costa Rica ...... 7 The Caribe Sur of Costa Rica ...... 8 Agriculture and conservation in Costa Rica ...... 9 Objectives, research questions, and predictions ...... 11

CHAPTER II METHODOLOGY ...... 15

Study Site ...... 15 Nonhuman primates of Gandoca, Costa Rica ...... 20 Los Gandoqueños ...... 22 Interviews ...... 23 Camera trapping and primate observations ...... 26 Data processing and analyses ...... 26

CHAPTER III RESULTS ...... 28

Animals identified as problematic and/or crop raiders ...... 28 Perceived behavioral differences between monkeys ...... 32 Feeding habits when raiding crops/gardens ...... 34 Perceptions and attitudes toward monkeys ...... 36 Management practices ...... 37 Importance of conservation ...... 40 Observations of feeding instances and camera traps ...... 42

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CHAPTER IV DISCUSSION ...... 45

Monkeys are among the most problematic animals in Gandoca ...... 45 Perceived differences between monkeys in Gandoca ...... 46 “Monkeys love the water apple” ...... 47 Perceptions and attitudes toward monkeys ...... 49 “Spider monkeys are my favorite monkey” ...... 53 Perceiving primates as human-like creatures ...... 54 Importance of wildlife conservation ...... 54 Management practices ...... 58 Recommendations ...... 62 Future directions ...... 65

REFERENCES ...... 67

APPENDIX A INSTITUTIONAL REVIEW BOARD APPROVAL LETTER...... 76

APPENDIX B PARTICIPANTS’ SOCIO-DEMOGRAPHIC DATA ...... 77

APPENDIX C PARTICIPANTS’ LAND INFORMATION ...... 78

APPENDIX D ADJECTIVES USED BY PARTICIPANTS TO DESCRIBE THE MONKEYS OF GANDOCA AND THEIR CLASSIFICATION, NUMBER OF RESPONSES AND PERCENT OF RESPONSES ...... 80

APPENDIX E ADJECTIVES USED TO DESCRIBE EACH SPECIES, THE CLASSIFICATION OF ADJECTIVES AND THE NUMBER AND PERCENT OF RESPONDENTS WHO USED EACH ADJECTIVE ...... 81

LIST OF FIGURES

Page

Figure 1 Map of Costa Rica with the location of some main towns in the Limón Province of the Caribe Sur ...... 17

Figure 2 Map of the RNMVS-GM and important location points ...... 19

Figure 3 Map of the areas interviewed in Gandoca and the approximate interview location points ...... 29

Figure 4 The percentage for each class of animals reported as problematic and/or crop raiders across 105 responses (N=24) ...... 30

Figure 5 The percent of responses that were positive, neutral, or negative for each monkey species...... 36

Figure 6 The respondent’s preference for the different monkey species ...... 37

Figure 7 Scare crow installed in the canopy of a rambutan tree to scare away raiders, only reported to work for oropendolas...... 39

Figure 8 The methods used by respondents to prevent or deter monkeys from entering their land and/or raiding their crops and gardens and the percent of respondents who used each method ...... 40

Figure 9 Percent of responses for the importance of conservation that had a utilitarian, intrinsic, and non-use value ...... 41

Figure 10 Yolillo fruit (Raphia taedigera) half-eaten by Ateles ...... 42

Figure 11 A black-handed spider monkey (A. geoffroyi ornatus) feeding on rambutan fruits (Nephelium lappaceum) ...... 43

Figure 12 Group of howler monkeys (A. palliata) resting near a banana (Musa spp) plantation ...... 44

Figure 13 The clearing of forests near their land is common to star new plantations as needed, such as for banana or plantain ...... 56

Figure 14 According to locals, several years ago, people from the Netherlands established a program where they gave monetary compensation to locals as incentive ...... 57

LIST OF TABLES

Page

Table 1 Studies reporting crop-raiding by the primates of interest in this study .... 5 Table 2 Estimated population density (individuals/km2) of A. palliata, A. geoffroyi, and C. capucinus in selected sites of Costa Rica ...... 8

Table 3 Formulated research questions for this study ...... 12 Table 4 Life history traits of primates of interest in this study ...... 21 Table 5 Range and mean percent time spent foraging fruits, leaves, flowers, and prey by primate species ...... 21

Table 6 Questions posed to the participants during the interviews ...... 24 Table 7 Animals reported by participants to be problematic and/or to raid gardens and/or crops ...... 31

Table 8 Respondents who reported Ateles, Cebus, and Alouatta as problematic and/or crop raiders and as most problematic ...... 33

Table 9 Food items reported by the participants to be consumed by the monkeys from their cultivated land or gardens ...... 34

Table 10 The number and percent of positive, neutral, and negative responses when describing the Gandoca monkeys ...... 36

Table 11 The number and percent of respondents for each mitigation methods against problem animals and crop/garden raiders and their level of perceived success ...... 38

Table 12 The reasons for wildlife conservation as stated by respondents ...... 41

NOMENCLATURE

NHP Non-Human Primates

RNMVS-GM Refugio Nacional Mixto de Vida Silvestre Gandoca-Manzanillo

YA Years Ago

UFC United Fruit Company

SINAC Sistema Nacional de Áreas de Conservación

FAO Food and Agriculture Organization

IRB Institutional Review Board

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ACKNOWLEDGEMENTS

Thanks to my parents, Gloria and Gilberto, for their unconditional support throughout my

“crazy idea” of leaving home to pursue my dream of studying wildlife. Thanks to my loving partner and friend, José, for putting up with me during this process and offering all kinds of support. Thanks to my major professor, Dr. Pruetz, for the opportunity of doing research under her guidance and for always finding a way to help me out. Also, my committee members, Dr.

Nuñez and Dr. Viatori, who were always willing to help by offering much needed advice and direction. Infinite thanks to Dr. Lindshield for being my mentor and friend during my undergraduate and graduate student years. I will forever carry all you advise with me. Without your help and guidance, I wouldn’t have accomplished as much. Thanks to my anthropology colleagues who were there whenever I needed to vent or simply have a good laugh. Finally, big thanks to the community of Gandoca who allowed me to conduct this study, especially the

Rodríguez Rojas family who took me in as another family member and Braulio Venegas Rojas for his assistance in the field.

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ABSTRACT

Crop-raiding by nonhuman primates is one of the most common of human-alloprimate interactions, often translating into conflict and reduced tolerance. Although it has not been reported to be as problematic in the Neotropics, examining this interaction is important for identifying potential conflicts and incorporating adequate management practices. In the community of Gandoca, Costa Rica, residents coexist with three sympatric primates: Alouatta palliata, Cebus imitator, and Ateles geoffroyi ornatus. We examined the attitudes and perceptions of 24 individuals towards alloprimates and how crop-raiding influenced them. We evaluated the intensity of such conflict as perceived by the community through semi-structured interviews. Although 75% of the respondents said that crop-raiding was not problematic, 23 animals were identified to raid crops and/or gardens. The three nonhuman primates were reported to raid gardens and crops, but Alouatta more so than Cebus and Ateles. Interestingly, they were all described as “wasteful” when feeding, and this behavior seemed to “bother” people more than the raiding itself. Moreover, Alouatta was more likely to be described negatively,

Cebus more neutrally, and Ateles more positively. In conclusion, raiding behaviors affect the residents’ attitudes and perceptions, but other behaviors and physical traits may also factor into people’s perceptions of monkeys.

CHAPTER I

INTRODUCTION AND LITERATURE REVIEW

This study explores the interactions between the residents of Gandoca, a rural community within the Refugio Nacional Mixto de Vida Silvestre Gandoca-Manzanillo in southeastern Costa

Rica, and three free-ranging primates: mantled howler monkeys (Alouatta palliata), white-faced capuchin monkeys (Cebus capucinus imitator), and black-handed spider monkeys (Ateles geoffroyi ornatus). Using an ethnoprimatological approach, I aimed to assess the perceptions and attitudes of humans towards alloprimates to evaluate crop raiding as a potential conflict. Previous studies have found human attitudes and perceptions to be affected by nonhuman primate behaviors positively and negatively, sometimes impacting conservation practices as a result.

Although Neotropical primates do not crop raid as frequently as Paleotropical primates, it is important to monitor this interaction and be cautious regarding New World monkeys as deforestation rates increase and human and nonhuman primate (NHP) niche expansion continues, consequently intensifying conflicts in the near future.

Ethnoprimatology

We are now living in the Anthropocene; a time in which humans are influencing change in ecologies at both local and global scales, faster than we can study it (Rose 2009). These ecological and cultural changes affect both humans and nonhuman primates. Therefore, anthropologists suggest to study all primates, human and nonhuman, as co-participants of a dynamic biological and cultural ecosystem (Fuentes and Wolfe, 2002). The emerging field of ethnoprimatology was first coined by Sponsel (1997), but many researchers in the 1990s

2 recognized the need to study human-nonhuman primate interactions given the increased intensity of rising conflicts. Nonetheless, this field examines relationships of coexistence and recognizes the cohabitation of primates before the dawn of anatomically modern humans. Ethnoprimatology is a hybrid field, which integrates primatological and anthropological methods. While anthropology is the study of ourselves, primatology is a subset of the field and crosses disciplines focusing on the study of nonhuman primates, our closest living relatives.

As we all aim to thrive in a changing environment, ethnoprimatological methods facilitate the assessment of conservation realities by considering both human and nonhuman primate mutual ecologies (Fuentes and Wolfe, 2002). Anthropologists are now more than ever including ethnoprimatological methods in their projects and it is critical that they do (Dolhinow

2002). A study conducted in Venezuela (Lizarralde 2002) investigated the Barí ecology, an indigenous group living in close relationship with alloprimates and how their culture and changing technologies impacted the populations of four monkey species. For example, Barí mythology includes the origin of monkeys and how their creator, Sabasebaa, told them they should eat monkeys. For the Barí, monkeys were a preferred food source over all other animals, monkeys were kept as pets, and their teeth are used for necklaces for the purpose of acquiring monkey characteristics. However, perceptions and attitudes towards the monkeys varied by species. For the Barí, spider monkeys (Ateles belzebuth hybridus) were the most precious of all.

These cultural preferences certainly impacted the way the Barí interacted with alloprimates

(Lizarralde 2002). Conversely, the way monkeys behaved might have influenced these initial perceptions. Finally, Lizarralde (2002) concluded that the increasing population density of the

Barí, the patterns of their settlements, and their uses of new technologies, such as firearms, are resulting in unsustainable resource use.

The human-nonhuman primate interface was described as one of the most important areas of anthropology in the 21st century (Fuentes and Hockings 2010) as primates, human and nonhuman, continue expanding their niches and overlapping. A more recent study (Ellwanger et al. 2015), conducted in Fanjingshan National Nature Reserve, in China, highlighted the importance of local people’s knowledge and attitudes towards the endangered Guizhou Snub-

Nosed monkey. For example, they found that respondents perceived conservation as a trade-off between costs and benefits to their livelihoods. Although people viewed the monkey positively, the survival of this species depends on the conservation of the forests. Ellwanger and colleagues

(2015) recommended the improvement of communication between residents and officials, implementing educational programs, and including the participation of the community in management decisions. In this study, understanding the “ethno” benefited the “primatology” goals. Currently, the goal of many primatological studies is to conserve and coexist with our closest living relatives. However, conserving culture is also significant (Fuentes and Wolfe

2002). Conservation requires action. People are critical for conservation and without involving people in conservation plans, these will not be successful.

Crop-raiding by nonhuman primates

Crop raiding is not a novel or rare occurrence. Crop raiding by wildlife has most likely been occurring since the beginning of human agricultural settlements more than 10,000 YA. The crop raiding phenomenon can be defined as the movement of wild animals onto agricultural land to feed on vegetables, fruits, cereals, and trees that humans grow for their own use and consumption (Sillero-Zubiri and Switzer 2001) .Many different taxa, including insects, birds and mammals are known to raid and damage crops and gardens, sometimes causing major crop loss

4 for farmers (Meligethes spp.: Rusch et al. 2013; birds and other vertebrates: Hill 1997;

Loxodonta africana: Hoare 2015; Elephas maximus:Webber et al. 2011; Cervus elaphus L. and

Dama dama L.: Wilson et al. 2009) . Once people’s livelihoods are threatened or perceived to be threatened, crop raiding is then considered a human-wildlife conflict.

Hockings and Humle (2009) define human-nonhuman primate conflict as an interaction that results in negative impacts to human social life, economy or culture, primate behavior, ecology, culture, or conservation. The crop raiding human-nonhuman primate conflict has been densely studied in the past few years, but most of the research is focused in Africa and Asia where NHP range near agricultural lands. The catarrhines are the best known crop-raiding primates, especially Papio, Chlorocebus (Hill 2005), and Macaca (Southwick et al. 2005).

However, other primates, Pan troglodytes (Reynolds 2005; Hockings 2009), Pan paniscus

(Hockings and Humle 2009), Gorilla beringei (Goldsmith et al. 2006), and Pongo abelii

(Campbell-Smith et al. 2010), also raid crops. American primates are thought to be less probable to raid crops in part because of their increased arboreality (McKinney et al., 2015). Nonetheless, a number of Neotropical primates have been reported to raid crops across Central and South

America (Table 1), with most reports coming from Costa Rica.

Across the world, nonhuman primates are the most frequently identified crop-raiding animals (Sillero-Zubiri and Switzer 2001). Altmann (1998) described most species of primates as eclectic omnivores. Eclectic omnivores exhibit in various degrees, dietary selectivity, flexibility and diversity. These characteristics are key for their ability to exploit new food sources and supplement their diets when something palatable or profitable is found (Altmann 2009) such as a banana crop, for example. In addition, their intelligence, social behavior and curiosity makes them particularly prone to raid crops and take advantage of anthropogenic habitats (Mckinney

2015). Moreover, the cooperative behaviors, manual dexterity, and communication skills exhibited by primates makes them particularly difficult to deter from human settlements. For example, they can quickly learn how to trespass electric fences (Strum 1994) and learn the time of the day or night when humans are not guarding croplands (Krief et al. 2014).

Table 1 Studies reporting crop-raiding by primates of interest in this study.

Genus Species Crop Country Source Ateles A. geoffroyi yucatanensis, pineapple, Belize, Waters and Ulloa 2007; A. geoffroyi ornatus banana Costa Rica Jill Pruetz pers. comm.

Alouatta A. palliata mango Costa Rica McKinney 2010 Cebus C. apella, potato, corn, Brazil, Galetti and Pedroni 2008; cassava, Costa Rica de Oliveira and de Souza C. capucinus imitator cacao, Fialho 2007;Baker and mango, Schutt 2005 musa spp.

A reliable predictor of crop raiding is the type of crops grown. Primates seem to prefer crops high in caloric content or simple sugars and that are easy to handle (Nijman and Nekaris

2010). Nonetheless, other factors, such as the distribution of the crop (Naughton-treves et al.

1998), habitat quality, forest food availability, and levels of human activity (Hockings et al.

2009) among other, also influence raiding frequency and intensity. Although crop raiding seems to be of significantly less common occurrence in the Neotropics than in the Paleotropics (Estrada

2006), it still remains as an important issue to address, and to assess how these behaviors affect human attitudes towards alloprimates.

Why study human attitudes and perceptions towards nonhuman primates?

An attitude is a mindset or tendency to act in a certain way due to both a person’s experiences and temperament (Allport 1937). Based on social psychological theory, attitudes arise out of general values, primitive beliefs and perceptions. Attitudes can be immediate predictors of behavior and are more likely to influence behaviors when they are strong, based on personal experiences, and salient (Pickens 2015). Perceptions are closely related to attitudes, and these are a process by which an organism interprets a stimulus into something meaningful based on prior experiences (Lindsay and Norman 1977). However, sometimes perceptions are substantially different from reality. In other words, studying how people feel about wildlife and how they perceive and relate to animals facilitates the understanding of human-nonhuman primate interactions. Previous studies have shown that cultural perceptions have an important role in shaping human interactions with the environment and with alloprimates (Lizarralde,

2002; Sousa et al. 2014; Riley et al. 2013). Moreover, conflicts may be identified this way and the development of mitigation strategies that are based on and target local people’s concerns

(Hill et al. 2002). An effective conservation program should be able to incorporate the local values regarding to resources being conserved, instead of focusing on the benefits as a conservationist may perceive it (Costa et al. 2013). For example, some people may find value in conservation if they can directly benefit at the individual level from the protection of wildlife and other resources.

A research study conducted in Northern Sumatra, Indonesia, explored local attitudes and perceptions toward crop-raiding orangutans (Pongo abelii) and other NHP (Campbell-Smith et al. 2010). In this study researchers were able to detect negative perceptions towards orangutans, that persisted regardless of farmers’ previous experience with crop raiding by orangutans.

Campbell-Smith and colleagues (2010) explained that most of these negative perceptions were driven by fear. In addition, there were local legends, as well as physical and behavioral traits that were impacting farmer’s perceptions and attitudes, therefore affecting how they responded to orangutan presence and crop raiding events. By using ethnoprimatological methods, they were able to detect factors driving negative perceptions and help educate the locals and encourage acceptance that orangutans are genuinely not dangerous. More importantly, they found that some people were tolerant of orangutans and supportive of their conservation even when experiencing or perceiving conflict. This study demonstrates the importance of involving the community in conservation efforts, learning more about the local perceptions and attitudes, and including their concerns in management and conservation plans.

Nonhuman primates of Costa Rica

There are four non-human primate species native to Costa Rica. The squirrel monkey (Saimiri oerstedii) is found only in the western region of Costa Rica and is currently listed as endangered by the IUCN Red List (Wong et al. 2008). Ateles geoffroyi ornatus, also listed as an endangered species by the IUCN Red List (Cuarón, Morales, et al. 2008), ranges through Nicaragua, Costa

Rica, and Panamá. Alouatta palliata palliata, a species of Least Concern (Cuarón, Shedden, et al.

2008), ranges from Mexico to northern Ecuador, with the largest geographic distribution of all.

Finally, the geographic distribution of Cebus capucinus imitator extends from Honduras to

Colombia, with no classification by the IUCN. Population size estimates of all three primate species for the whole of Costa Rica are unknown, but data on population density estimates within some protected areas is available (Table 2), not including RNMVS-GM (DiFiore, Link, and

Campbell 2011). Overall, howler monkeys are often described as the most common species of

8 the country, followed by capuchins, spider monkeys, and squirrel monkeys (Zaldívar et al. 2004).

However, this might vary by region as shown in Table 2. The apparent difference in population density could be due to differences in life history traits (Table 4). For example, while A. palliata and C. capucinus show a higher reproductive rate, A. geoffroyi has a relatively slow one (Fedigan and Jack 2001).

Table 2 Estimated population density (individuals/km2) of A. palliata, A. geoffroyi, and C. capucinus in selected sites of Costa Rica.

Population density (ind/km2) Study site Alouatta palliata Ateles geoffroyi Cebus capucinus El Zota Biological Field Station1 8.3-8.5 12-12.5 5.9-6.0 Santa Rosa National Park2 7.9 7.9 7.5 Palo Verde3 59.3 0.62 15.4 1Lindshield 2006; 2Sorensen and Fedigan 2000; 3Massey 1987

The Caribe Sur of Costa Rica

The area of the Caribe Sur1 (Figure 1) was first inhabited by indigenous people who preferred to live further from the mountains and closer to the coast. Around the 1750’s, the first group of turtle hunters and fishermen arrived (García and Grant 2012). This group of turtle hunters and fishermen came from Nicaragua and Panamá, and are said to be descendants of the

Miskitos native group, and Afrocaribbeans who traveled from Jamaica to Central America. They sailed in small boats and camped on the coast from March to September, during the turtle nesting season. During their stay in the Caribe Sur coast, they planted yucca, plantain, yam and coconut,

1 The Caribe Sur region refers to the southeastern area of Costa Rica adjacent to the Caribbean Sea.

9 among other. They also started planning for their next season, planting crops to harvest on their next visit. In these small camps, they built shelters using the resources they found, such as chonta

(Socratea exorrhiza) and yolillo palm (Raphia taedigera) leaves. Finally, in 1828, several groups of turtle hunters and fishermen settled with their families in the Caribe Sur coast of Costa Rica.

During the beginnings of their settlement, their work was mostly dependent on turtles.

The shell of hawksbill turtles (Eretmochelys imbricata), was removed and sold to companies that would send those to Germany and were used for the production of buttons and combs (García and Grant 2012). Also, green sea turtles (Chelonia mydas) were hunted for their meat and oils.

The first inhabitants relied on fishing and hunting to survive and one of the most important cultivated products was the coconut palm, followed by yucca. There were not many fruits here, and many were brought from the San Andrés island, including also domestic animals. Other fruits, such as the mango, were brought from Jamaica.

Later, the United Fruit Company (UFC) set foot in Costa Rica and started constructing the railroad in 1908. This move changed the landscape across the Talamanca mountains as the land turned into banana plantations along the railroad. During those times, the inhabitants of the coast would enter the mountains and sell or trade their fishery products to North Americans working for the UFC or indigenous groups. At the turn of the century, agriculture grew stronger in the coast with the arrival of more people from Nicaragua and Panamá, while Jamaicans cultivated cacao inland. Other crops, such as pineapple, watermelon, sugar cane, cucumber, tomato, okra, and rice, were destined for consumption.

Agriculture and Conservation in Costa Rica

According to the 2014 Food and Agriculture Organization (FAO) report,2 nearly 35% of

Costa Rica’s land is used for agricultural purposes, that is, land that is arable, under permanent crops, and under permanent pastures. Nonetheless, 8,000 hectares are plantations with no chemical treatments, and over 3,000 farmers offer organic certified products for the national and international market. In 2009, PROCOMER (La Promotora del Comercio Exterior de Costa

Rica) reported more than 36,000 tons of organic products to be exported, with a total value of more than 26 million dollars (Chaves et al. 2010). Most of the exports from Costa Rica go to

Europe, United States, Canada, Australia, and Japan. The Talamanca canton is one of the main areas with organic farms, as well as the Zarcero, Turrialba, Zona Norte and Cartago zones

(López Navas and Soler Zurita 2014).

Costa Rica is the seventh largest producer of bananas in the world. Bananas, as well as other fruits, are the main agricultural export and these are cultivated across the Great Central

Valley (Gran Valle Central), the Northern Zone (Zona Norte), and the Pacific and Caribbean coast. Roughly 8% of Costa Rica’s gross domestic product (GDP) is made up of agricultural exports. Coffee, also known as the golden bean, is another major crop produced in Costa Rica, and it was the motor of the economy during the 19th century. The golden bean is still the single most important crop in terms of land use and production value, followed by rice. Sugar cane, pineapple, and cacao are also major crops important for Costa Rican the economy.

2http://data.worldbank.org/indicator/AG.LND.AGRI.ZS?end=2014&locations=CR&start=1961&view=ch art

One of seven Costa Ricans work in the agricultural sector (López Navas and Soler Zurita

2014). In this small Central American country, agriculture is the main source of income for the population of lower level of education, whereas most people with secondary education mainly work on tourism (IICA 2010). Approximately 40% of the rural population over 15 years of age, work in agriculture related jobs, more so in the Limón Province (INEC 2012). Bouroncle and colleagues (2015), explained that the more dependent the population is on agriculture, the more they will be impacted by climate change in the coming years.

Although agriculture and ranching is one of the most important sectors in the Costa Rican economy, tourism is actually the main economic activity (López Navas and Soler Zurita, 2014).

Conservation efforts in Costa Rica have been recognized worldwide, and their national parks and protected areas have brought people from all around the world to contemplate and enjoy of many different touristic activities. Moreover, nearly 20% of the land has been declared World Heritage sites by UNESCO. All of the resources put into conservation has made Costa Rica one of the most popular destinations for ecotourism and the first ranked nation among Latin-American countries in their environmental performance index (EPI) as of 2016 (Hsu et al. 2016).

Objectives, research questions, and predictions

Objectives

In July of 2015, I had an informal conversation with a Gandoca resident. The local woman explained how all three species of monkeys could be easily observed in the community.

Further, she explained how monkeys entered her garden and fed from the fruits and crops they were growing for the consumption of her family:

“All three species of monkeys can be seen around here. They frequently come into my garden and they eat everything. They bite the fruit and throw it away, both ripe and unripe. At least the spider monkey is more selective, but the howler is lazy and very wasteful. The capuchins are bandits. Sometimes one has to keep track of what fruit is about to ripen, so that you pick it before animals come and ruin it.”

This conversation with the Gandoca resident sparked my interest in learning more regarding the human-nonhuman primate interactions in Gandoca, Costa Rica. Therefore, the main objectives of this study were to explore the human-alloprimate interface in the community of Gandoca, Costa Rica and how the attitudes and perceptions of the people were affected by the monkeys’ behaviors. More specifically, my goal was to examine how crop raiding by monkeys influenced attitudes and perceptions of the residents of Gandoca towards these animals. I aimed

Table 3 Formulated research questions for this study.

Is crop raiding by primates perceived as a problem in Gandoca?

How differently are the local people perceiving the three monkey species? Is one monkey species perceived as more problematic than others?

How are crop raiding behaviors affecting the local people’s attitudes?

Are there any demographic factors determining attitudes?

Are there any mitigation methods to deter or prevent monkeys and other animals from feeding from crops and gardens?

to evaluate the intensity of such conflicts as perceived by the community through semi-structured interviews of the local land owners and farmers. In addition, I intended to examine whether they perceived any behavioral differences between the three primate species and if these were

13 creating different attitudes. Finally, I wanted to observe and record monkeys feeding behaviors through vigils and camera traps in order to confirm the locals’ reports and better evaluate this interaction. From these objectives I developed five main research questions (Table 3). I will explain how these objectives were accomplished and how these questions were addressed in detail in the methods section (Chapter 2).

Predictions

I predicted the people of the Gandoca community would consider crop raiding a problem, and monkeys to be problematic raiders. Based on the informal conversation with the local woman, I expected similar attitudes and perceptions throughout the community about monkeys.

However, I predicted that residents relying more heavily on agriculture would have a more negative attitude towards crop and garden raiding primates. Moreover, I expected different perceptions and attitudes towards the different primates based on perceived behaviors. I predicted that there would be a relationship between crop raiding frequency and intensity, as reported by participants, and their attitudes. Therefore, the more a monkey is reported to crop raid, the more negatively it is going to be perceived.

Mantled howler monkeys, while considered part of the most folivorous primate taxon in the

New World (Milton 1980), they have an incredibly flexible diet ( Chapman 1987; Arroyo-

Rodríguez and Dias 2010), can have small home ranges and possess energy saving strategies that are key for their ability to persist in spite of extreme anthropogenic disturbance to their habitats

(Katharine Milton 1998; Korstjens et al. 2010). These characteristics are known to allow howler monkeys to survive in matrix habitats better than other New World monkeys and maintain a widespread distribution (McKinney et al. 2015). Due to their prevalent distribution, adaptability to disturbed habitats, and more conspicuous behaviors, I expected howler monkeys to be

14 described as the most problematic species among the three species of monkeys in Gandoca, even though the other species are considered more frugivorous.

Capuchin monkeys have been previously reported as crop raiders in different countries across Central and South America (Galetti and Pedroni 2008; de Oliveira and de Souza Fialho

2007; Baker and Schutt 2005). This is probably given to their increased terrestrial locomotion and foraging in comparison to other American primates, as well as their more curious behaviors.

Capuchins have been described as “destructive” foragers (Terborgh 1983) because when they search for foods they move through different surfaces, uncurling dried up leaves, shifting things around, breaking things apart and peeling back coverings from trees and fruits (Panger et al.

2002). They are also known to be aggressive towards humans in other regions of Costa Rica when trying to steal food from park visitors (Campbell 2013). For all these reasons, I expected the white-faced capuchin monkeys in Gandoca to be a problematic crop and garden raider as well, and to be perceived negatively. However, this species did not seem to be as widely distributed in Gandoca, in comparison to howler monkeys

Finally, I predicted spider monkeys to be perceived more positively. Although spider monkeys are highly frugivorous and considered to be ripe fruit specialists (Colin A. Chapman

1990), I predicted that this species would not be reported to raid gardens and crops as frequently as howler and capuchin monkeys. Based on previous observations and communication with residents, spider monkeys are not seen as frequently as the two other primate species in the forest edge of the Caribe Sur region. Also, not many studies have reported spider monkeys to raid crops or gardens, although they have been observed in agroecosystems before (Estrada and Coates-

Estrada 1996).

CHAPTER II

METHODOLOGY

This study was conducted in the rural community of Gandoca, located within and adjacent to the Refugio Nacional Mixto de Vida Silvestre Gandoca-Manzanillo, Jairo Mora

Sandoval, in southeastern Costa Rica. Semi-structured and open-ended interviews were conducted to explore human attitudes and assess the interactions between residents and the three nonhuman primates. Participation was completely voluntary, and no compensation was offered.

Participants were asked a variety of questions, ranging from socio-demographic information, attitudinal data, observations about primate behavior and ecology, crop/garden raiding by wildlife and management practices to mitigate such behavior (Table 6). In addition, camera traps were installed in locations of reported crop and garden raiding to capture and confirm the consumption of food items. Data were analyzed using JMP Pro 12 statistical software. I used descriptive statistics to compare variables, and contingency analyses were used to identify associations between socio-demographic variables, attitudinal data, and other variables.

Study Site

The Refugio Nacional Mixto de Vida Silvestre Gandoca-Manzanillo, Jairo Mora Sandoval

The study was conducted at the Gandoca sector of the Gandoca-Manzanillo national wildlife refuge in southeastern Costa Rica (9°35'41.7"N 82°36'20.8"W) from June through July of 2016 (Figure 2). The wildlife refuge is part of the Corredor Biológico Talamanca Caribe and encompasses a total of 9,449 hectares. Of those, 5,013 hectares comprise the terrestrial portion of the refuge and 4,436 comprise the marine portion (Fonseca Borrás et al. 2005). The Refugio

Nacional Mixto de Vida Silvestre Gandoca-Manzanillo (RNMVS-GM) is located in the

Talamanca canton of the Limón Province, in the Sixaola district, and it pertains to the Area de

Conservación La Amistad del Caribe (ACLAC). The refuge was formed in July 1985 through an executive order by Presidential Decree 16614-MAG01 in an effort to protect the natural resources of the region. A decade later, the executive decree 25595-MINAE established the base for the co-administration of a protected area. Then, a committee formed by the MINAE (Ministry of the Environment and Energy), local associations, non-governmental organizations, and the municipality of Talamanca, was created to manage the refuge. As the name of the RNMVS-GM indicates, the refuge is of mixed type, which means that its land is owned not only by the state, but also by private owners (90% is privately owned). The agreement was to protect the area, while also protecting the right of the people to remain in their land. As a result of such agreement, the sustainable management and conservation of the refuge is carried out in parallel with the participation of the communities, as previously mentioned. In addition, this means that people cannot continue expanding their land through deforestation (García and Grant 2012).

Later, in 2013, the refuge was renamed Refugio Nacional Mixto de Vida Silvestre Gandoca-

Manzanillo, Jairo Mora Sandoval, in honor of the turtle conservationist born and raised in

Gandoca. Jairo was murdered in May of 2013 at the beach of Moín, north of the refuge, after patrolling that same beach with volunteers to protect sea turtle nests from poachers (Fendt 2015).

The Gandoca-Manzanillo community is known as an active civil society, which has successfully fought against the overuse of agrochemicals, banana labor exploitation, and transnational oil exploration (Fonseca Borrás et al. 2005). The active participation of the community within protected areas is a vital part of the National System of Conservation Areas

(SINAC) in Costa Rica. Local councils were established within communities near conservation areas with the

Caribe Sur

Figure 1 Map of Costa Rica with the location of some main towns in the Limón Province (white) of the Caribe Sur.

executive decree 22481-MIRENEM in 1993. This was later reinforced by the National Plan of

Development 1994-1998, which considers the need to reinforce civil participation and have them assume their role as active individuals of development. These councils would work as bodies of coordination, support, and the discussion of conservation and sustainable development programs.

For example, the Asociación Microempresarial de Productores(as) Agropecuarios de Gandoca

(ASOMIPAG) is an association directed by the residents and for the residents. ASOMIPAG works for the sustainable development of the community through ecotourism, volunteer

18 programs, and other projects that benefit the members of the community while preserving the natural beauty of Gandoca.

The Costa Rican Caribbean southern coast, also known as Caribe Sur (Figure 1), is characterized by its biological richness, as well as its cultural richness. Today, RNMVS-GM is comprised of two communities with marked social and economic differences. The community of

Manzanillo is characterized by an important touristic development, a predominant Afro-

Caribbean culture and a high volume of European immigrants. On the other hand, the community of Gandoca, which is my population of interest, is mostly composed of locals with Afro-

Caribbean and indigenous descent, that dedicate their life to banana plantations, agricultural work, forestry, ecotourism and scientific tourism (Weitzner and Fonseca 2000).

Environment at Gandoca

The geomorphology of RNMVS-GM is characterized by coast, coastal plain and mountainous areas. The weather reflects the general pattern typical of the Costa Rican southeastern Caribbean, distinguished by the movements of subtropical anticyclones of the

Atlantic and North Pacific, known as the trade winds. Precipitation occurs throughout the year, ranging from 1950 to 3000 mm annually, with a decrease in March-April and September-

October. The average annual temperature is between 24o C and 27o C (Fonseca Borrás et al.

2005; García and Grant 2012). The climate in the refuge area allows for the establishment of a great variety of soil uses, from forests to diverse annual crop plantations.

The RNMVS-GM is classified as a tropical humid forest consisting of remnants of rainforest, coastal lagoons, palm swamps, swamp forest and wooded areas. In addition, RNMVS-

GM holds freshwater and marine habitats that include the most important coral reef of the Costa

Rican Atlantic coast, a freshwater swamp and the only mangrove swamp of the Atlantic coast of

19 the country (Dirección General de Vida Silvestre 1988;Fonseca Borrás et al. 2005). The vegetation consists mainly of relatively tall forests of approximately 30 to 40 meters of height and generally of three strata. Epiphytes are abundant, as well as species such as Dipterix panamensis, a tropical

Figure 2 Map of the Refugio Nacional Mixto de Vida Silvestre Gandoca-Manzanillo, Jairo Mora Sandoval and important location points.

forest emergent species, and Prioria copaifera, as well as a variety of Bromeliads (Aechmea,

Cryosophila), among others. The forests are important habitat for the endangered black-handed spider monkey (Ateles geoffroyi ornatus), and the coastal region serves as important nesting sites

20 for three species of sea turtles: the endangered green turtle (Chelonia mydas), the critically endangered hawksbill turtle (Eretmochelys imbricate) and the leather-back sea turtle

(Dermochelys coriacea). For all of these reasons, RNMVS-GM has great potential for ecotourism and scientific tourism.

Non-human primates of Gandoca, Costa Rica

The southeastern region of Costa Rica hosts three species of NHP: the black-handed spider monkey (Ateles geoffroyi ornatus), the mantled howler monkey (Alouatta palliata palliata), and the white-faced capuchin monkey (Cebus capucinus imitator). The three species occupy a variety of altitudinal ranges and habitats, including mangroves, riparian forests, dry and humid forests, lowland, premontane, and montane forests, secondary forests, and forest edges

(Eisenberg 1983; Freese 1983; Glander 1983; Sánchez 1991; Stoner 1996; Carrillo et al. 1999;

Mora 2000; Clarke et al. 2002). Although all three species use a wide variety of habitats, they differ in habitat preference, diet, home range, size and other characteristics (Table 4 and Table

5). For example, Ateles is restricted to large forest fragments and tends to be less common in disturbed or altered environments (Sorensen and Fedigan 2000; Zaldívar et al. 2004). On the other hand, Cebus and Alouatta can occupy small patches of forests and are more commonly found in disturbed habitats (Carrillo et al. 1999). These species also vary in the proportion of time spent eating fruits, flowers, leaves, or insects. As shown in Table 5, Ateles and Cebus spend more time foraging for fruits, while Alouatta spends a higher proportion of time foraging for leaves.

As previously mentioned, all three primate species found in Gandoca have previously been reported to crop raid at other locations (Table 1). The rapid conversion of tropical forests into

21 agricultural land has caused incredible habitat loss and fragmentation for NHP. But, not all species decline toward extinction following fragmentation; some primates might demonstrate resilience in matrix habitats (Estrada et al. 2005).

Table 4 Life history traits of primates of interest in this study. Adapted from Zaldívar et al. 2004.

Ateles geoffroyi Alouatta palliata Cebus capucinus Average male size (gr) 8375 6528 3333 Average female size (gr) 6624 4020 2283 Age at first birth (yr) 5.62 3.58 4.00 Interbirth interval (yr) 2.66 1.88 1.60 Birth rate 0.38 0.59 0.63 Population growth rate 0.113 0.178 0.172 Home range (ha) 25-98 27-91 50-100 Average home range (ha) 2991* 291* 802 Average day range (meters) 1,8851 4871 - 1 2 Di Fiore, Link, and Campbell 2011; Freese and Oppenheimer 1981 *Group or community range in hectares

Table 5 Range and mean percent time spent foraging fruits, leaves, flowers, and insects by primate species. Adapted from Zaldivar et al. 2014 Range (and mean) percent time foraging for:

Species Fruits Leaves Flowers Insects/Prey Sources

Ateles geoffroyi 71-78 13-11 14-10 1-2 Chapman 1987; Chapman and Chapman 1991; Chapman, (71.4) (12.5) (14) (2.1) Chapman, and Wrangham 1995)

Alouatta palliata 13-29 49-64 18-23 0 Chapman, 1987; Glander, 1981; Stoner, 1996 (23) (68) (8.5) (0)

Cebus capucinus 53-81 1-15 0-2 17-45 Chapman, 1987; Chapman and Fedigan 1990 - - - -

Los Gandoqueños

Los gandoqueños, the people of Gandoca, are part of a small community located in the

Caribe Sur of Costa Rica, within the Refugio Nacional Mixto de Vida Silvestre Gandoca-

Manzanillo (RNMVS-GM). The area of the refuge has a low population density of 23 habitants per squared kilometer (García and Grant 2012), mostly concentrated in Punta Uva, Manzanillo,

Gandoca and San Miguel (Figure 2). Their most important resource is their soil, in addition to the beach and the lagoon. They rely mainly on agriculture for subsistence, but tourists are an important source of income for some. About half of the community benefits from ecotourism, these being those who are the closest to the beach and the lagoon that own cabinas3 and offer guided tours. Although the two sectors are next to each other, Gandoca is not as accessible as

Manzanillo, as there is no road that connects Gandoca to Manzanillo. The only way for visitors to get to Gandoca is to travel to Sixaola, the city on the border with Panama, and take a taxi or use some type of private transportation to access the road to Gandoca (Figure 2). In addition, the road to Gandoca is not paved, making it an even longer drive. This lack of access from

Manzanillo has made Gandoca and Manzanillo into very different communities economically and culturally.

In the Manzanillo community, a greater economic development stands out in comparison to Gandoca. Given that Manzanillo is more accessible, it receives a larger wave of tourists and foreigners while Gandoca receives only a few very brave and determined visitors. In Manzanillo, there are numerous restaurants, bars, and small supermarkets that serve many local and foreign visitors. Also, there is a greater number of lodging facilities with more elaborate buildings that

3 cabina: cabin/room for rent

23 include air conditioning, internet access, television, and other commodities. The residents of

Manzanillo economically depend on tourism, through lodging businesses and guided tours that include hiking, adventure, and fishing activities. In addition, members of the Manzanillo community work as artisanal fishers, street vendors, and state employees, among other employment forms. However, many of these local businesses are owned by foreigners (García and Grant 2012).

In contrast, commerce is not as developed in Gandoca. Most of the tourists visit Gandoca during the sea turtle nesting season, from February to August, and many students visit from national universities to volunteer or learn about the refuge and the Gandoca community. There are few cabinas in this sector and most are close to the beach and the lagoon where people can participate of touristic activities. The residents who do not own a business work on ranches or the banana plantations near Sixaola. The Gandoca community only has a few small stores with limited supplies and what they cannot get from their land, they get from markets found in

Sixaola, Bri-Bri, Hone Creek, or other towns. Los gandoqueños have had to learn how to be self- sufficient, given that they are disconnected from Manzanillo and other main cities of the area.

They have resorted to cultivating for their families, raising animals for meat, eggs, and milk, fishing, and sometimes even hunting for their subsistence.

Interviews

Door-to-door interviews were conducted in the sector of Gandoca within and adjacent to the

RNMVS-GM. Residents were asked to voluntarily participate, individually, without monetary compensation in a semi-structured interview in which questions were open-ended (Table 6). I

Table 6 Questions posed to the participants during the interviews.

Questions posed to the local community members of Gandoca, CR. Adapted from Chalise and Johnson, 2005, and Srivastava and Begum, 2005 with modifications by G.N.R

Respondent’s age and household holdings

1. What is your age and gender? 2. How many people are there in your household? 3. For how long have you been living in Gandoca? 4. How much land do you have under cultivation approximately? 5. How many crops are cultivated in your land, approximately? Which are these? 6. How far away is your crop land from the forest (approx.)?

Information about ranging monkeys

7. Is crop raiding an issue for you? 8. What animals do you see raiding your crops and/or gardens? 9. Which one is the most problematic? 10. Which of the three species visits your land? How frequent? 11. When monkeys visit your land, do they crop-raid? 12. What crops are the animals feeding from or damaging?

Attitude towards monkeys and wildlife conservation

13. Have you observed any marked differences between crop-raiding behaviors among the three monkey species? 14. Which of the three species raids crops more frequently? 15. Which monkey species causes the most damage or loss to your crop land? 16. How important is wildlife conservation? 17. Which species do you like the least and/or the most? 18. How would you describe the relationship between animals and the Gandoca community? How about with monkeys?

Management practices

19. Do you practice any specific methods to deter, protect or prevent monkeys from crop raiding? 20. How successful have been those methods? 21. Would you like to learn ways to manage and conserve the monkeys at the same time?

25 collected socio-demographic data, information on farmland and crops cultivated and human attitudes and behavioral data. The houses near the main road of Gandoca were surveyed, as well as houses in adjacent roads. The main road of Gandoca extends from Mata de Limón to Playa

Gandoca for approximately three kilometers where I opportunistically talked to people going door to door. In addition, I used the snow ball sampling method, in which I asked participants at the end of their interview about other potential participants with cultivated land.

I decided to have open-ended questions in order to get more information and explore additional themes and issues within the community regarding human interactions with wildlife and conservation, among other topics. Crop raiding was defined as any instance of one or more animal coming into the participant’s land and feeding from the fruits, vegetables, cereals, or any product grown for human consumption. Questions about crop raiding started broadly and became more specific to monkeys to avoid any influences from the interviewer or information loss about other animals seen crop raiding.

People were not directly asked about their attitudes or feelings because this may cause them to think I was expecting something specific. I wanted the conversations to flow more organically and most of them shared descriptions when I asked about any perceived differences between the alloprimates, which included different adjectives. These adjectives were then classified as negative, neutral, or positive.

Institutional Review Board (IRB) approval was requested and approved on May 23, 2016

(See Appendix A for letter of approval).

Camera trapping and primate observations

In order to better study the interactions between humans and monkeys in Gandoca and gain a better understanding from the primate perspective, I installed camera traps and conducted primate observations. After interviews were conducted, landowners were asked for permission to install camera traps in areas where potential raiding would happen. In addition, I asked for permission to conduct primate observations or vigils alongside their plantations within their privately-owned land. I installed three Stealth-Cam P18 7 Megapixel Compact Scouting Camera each in two different areas. These were placed in three different locations within the selected area, each at a different angle towards fruiting trees identified by landowners to be raided by monkeys. Cameras were inspected daily for new footage.

I intended to conduct vigils in areas where primates are known to raid. However, landowners indicated that this rarely happened and that monkeys would refrain from raiding with human presence. Therefore, I asked for permission and ventured into the farmer’s land and areas adjacent to their land to search for monkeys and record what they were feeding on. These areas adjacent to plantations included agroecosystems, mangrove swamps, and secondary forests.

Data processing and analyses

Interview data were transcribed from recordings or notes and coded when appropriate.

Responses were coded and aggregated by theme and percentage data were generated from the coded information. I used grounded theory to analyze open-ended responses after transcribed, in order to identify any important emerging themes (H. R. Bernard and Ryan 1998). When multiple responses were given to a question, data were presented as the percentage of respondents giving each response, and so may sum to over 100% (Gillingham and Lee 1999). Other percentages

27 were calculated across the number of responses instead of the number of respondents. To analyze the data, I used a contingency analysis, also known as the Pearson’s chi-square test, to assess the relationship between socio-demographic data and responses given by participants to questions about monkeys and management practices.

Because participants were not directly asked about their attitudes or perceptions, these were classified by examining the way people talked about monkeys. To do this, I went through the transcribed interviews in search of adjectives referring to monkeys. I coded each adjective as positive, neutral, or negative (See Appendix D). Perceptions of monkeys were then classified similarly, based on these adjectives. Because each individual participant had different perceptions for each species, it was difficult to determine their overall perception of monkeys, and I focused on their perception towards each species. Perception data were presented as response frequencies for the entire sample (Bauer 2003; Nekaris, Boulton, and Nijman 2013). I used the software package JMP Pro 12 to perform all statistical analyses.

CHAPTER III

RESULTS

From June 6 to July 31 of 2016, a total of 24 interviews were conducted in the community of Gandoca, Costa Rica (See appendices B and C). Participants comprised a total of

24 respondents: 12 men (50%) and 12 women (50%). The mean household size was 3 people

(range: 1-5). Nearly a third of the population (35%) fell between the ages of 31 to 40 years old.

More than half of the sample population (63%) was born and raised in Gandoca, while the rest moved to Gandoca as children or adults. All participants reported to have an area of cultivated land, including plantations and/or gardens with fruit trees, with an average area of 4.95 hectares

(range: 0.75-40). Over 70% of the people interviewed reported to cultivate both for sale and for their own consumption. The community was divided into the regions of Playa, Pueblo and

Bonifé (Figure 3). The majority of the people interviewed were from the Pueblo region (46%), followed by the Playa (25%) and Bonifé (29%) regions. Only 29% of the respondents directly benefit from tourism, either by owning cabinas, pulperías4 or sodas5, and/or by offering guided tours, as well as other services such as surf lessons. Of the eight participants who directly benefit from tourism, 86% live in the Playa region.

Animals identified as problematic and/or crop raiders

When participants (N=24) were asked if they considered crop raiding to be problematic, only 25% said yes, and 75% said no. A contingency analysis was performed to examine whether

4 pulpería: a store that sells groceries, usually small 5 soda: in Costa Rica, it is an establishment where food is prepared and served, usually small and local

Figure 3 Map of the areas interviewed of Playa, Pueblo and Bonifé and the approximate interview location points.

30 there was an association between area of residence and reporting crop raiding as a problem.

Although no association was found, (X2[2, N=24] =5.01, p=0.0852), people living in the Pueblo area tended to describe crop raiding as a problem more frequently (45.5%) in comparison to the participants living in the Playa (16.7%) and Bonifé (0%) regions. Participants identified a total of

23 animals from four different classes (across 105 responses) to be problematic in general or harmful to crops and/or gardens’ fruits (Table 7). Mammals were more commonly reported as problematic or crop raiders (61%), followed by birds (35%), insects (3%), and reptiles (1%)

(Figure 4). Across the 105 responses of crop raiding and problematic animals, 31% were birds,

30% were primates, 17% were mammals from the Order Carnivora and Didelphimorphia, and

14% were mammals from the Order Rodentia. The least common animals to be reported as problematic or raiders were insects (4%), reptiles (2%) and the opossum (1%). Of all the birds mentioned by respondents, the Montezuma oropendola (Psarocolius montezuma) and birds from the Psittacidae family were the most frequently mentioned.

Reptiles Insects 1% 3%

Birds 35%

Mammals 61%

Figure 4 The percentage for each class of animals reported as problematic and/or crop raiders across 105 responses (N=24).

Table 7 Animals reported by participants (N=24) to be problematic and/or raid gardens and/or crops.

Spanish name English name Scientific name IUCN No. Percentage Status respondents of respondents Birds Carpintero Woodpecker Picidae family - 3 13 31% Chachalaca Gray-headed chachalaca Ortalis cinereiceps LC 1 4 Loras y Pericos Parrots and Parakeets Psittacidae family - 11 46 Oropéndola Oropendola Psarocolius Montezuma LC 8 33 Pava Crested guan Penelope purpurascens LC 4 17 Tucán Keeled-billed toucan Ramphastos sulfuratus LC 2 8 Black-mandible Ramphastos ambiguus, NT Collared aracari Pteroglossus torquatus LC Other birds various species - 4 17 Primates Cariblanco, Carilla White-faced capuchin Cebus capucinus imitator ND 11 46

30% monkey 31

Colorado, Araña Black-handed spider Ateles geoffroyi ornatus EN 5 21 monkey Congo, Aullador Mantled howler monkey Alouatta palliata palliata LC 15 63 Other Gato de monte Tayra Eira barbara LC 5 24 mammals Yaguarundi Jaguarundi Herpailurus yagouaroundi LC 1 4 17% Kinkajou Kinkajou Potos flavus LC 1 4 Mapache Racoon Procyon lotor and/or LC 3 13 Procyon cancrivorus Manigordo Ocelot Leopardus pardalis LC 1 4 Pizote Coati Nasua narica LC 6 25 Zorro Opossum various species - 1 4 Rodents Ardilla Squirrel Sciurus variegatoides LC 12 50 14% Sciurus granatensis Guatuza Agouti Dasyprocta punctata LC 2 8 Tepezcuintle Paca Cuniculus paca LC 1 4 Insects 4% Insectos Insects various species - 4 17 Reptiles 1% Boa Boa constrictor Boa constrictor imperatus ND 2 8

The species that was most cited by respondents as problematic and damaging to crops and gardens was Alouatta palliata palliata (63% of respondents). Immediately following was

Sciurus spp. (50%), and Cebus capucinus imitator (46%). Predators, such as the Boa constrictor,

Eira barbara, Herpailurus yagouaroundi, Leopardus pardalis, and the gavilán (Fam.

Accipitridae), were problematic because they preyed on poultry. However, E. barbara also ate fruits such as Quararibea cordata. Only one respondent declared not to have seen any animals raiding crops or gardens.

When participants were asked to name all animals seen crop raiding or eating from their gardens, 71% reported to have seen at least one monkey species. Over half of the respondents had seen A. p. palliata (63%) eating from their crops and/or gardens, followed by C.c. imitator

(46%) and A.g. ornatus (21%). However, 60% of the respondents who mentioned Ateles as a problematic crop and/or garden raider, also described them as the most problematic of all the animals who visited their land.

Table 8 Respondents who reported Ateles, Cebus, and Alouatta as problematic animals and/or crop raiders.

Monkey species No. respondents No. respondents who reported to who reported as most raid/damage gardens/crops problematic monkey (%) (%) Ateles geoffroyi ornatus 5 (21) 3 (60) Cebus capucinus imitator 11 (46) 4 (36) Alouatta palliata palliata 15 (63) 5 (33)

Perceived behavioral differences between monkeys

Overall, people perceived behavioral differences between the three species of monkeys.

Seven participants (29%, N=24) reported to have seen no behavioral differences between

33 monkeys and/or not to pay attention. Two other respondents (8%) said that all three species were similar in behavior and equally destructive and wasteful when feeding but still described what they saw. The majority of the participants (63%) reported to have observed some behavioral differences and described those. Overall, all monkey species were described as harmful and wasteful when eating (n=16 responses), but Alouatta (50%) more so than Ateles (25%) and

Cebus (25%).

In general, Alouatta was more commonly seen visiting the participants’ land. Although they were the most frequently reported as a problem animal and/or crop raider (N=24, 63%) in comparison to the other two monkey species, it was not as frequently considered the most problematic of all animals (Table 8). In general, people described Alouatta as calm, friendly with people, and tame but also very harmful to their crops and/or gardens and wasteful when feeding.

People also explained that Alouatta stays for longer periods of time when visiting their land, maybe causing more harm not only by eating from their land, but also by bringing filth and bad odors. In contrast, Ateles was described as a more agile monkey, pretty, but still shy or untamed

(arisco). Most people explained that Ateles visits, takes what it wants to eat and leaves, and depending on the fruit, they would come back every day for more. They were still reported as wasteful and harmful to crops but at a lower frequency (21%) compared to Alouatta (Table 8).

On the other hand, Cebus was reported to be seen very rarely and was mostly described as untamed or shy (arisco). Cebus was the least described species in terms of their habits but was still the second most frequent monkey cited as problematic and/or crop raider by participants

(46%) and overall the third most problematic animal (Table 7).

Feeding habits when raiding crops/gardens

Participants were also asked to specify what monkeys were eating from their cultivated

lands or gardens. Respondents listed a total of 17 food items (across 78 responses) including

crops, wild species, and nonnative fruit trees (Table 9). The most cited food items were Syzygium

malaccensis (75%) and Nephelium lappaceum (46%), followed by Chrysophyllum cainito and

Psidium guajava (38% of respondents), all of which were eaten by the three species as reported

Table 9 Food items reported by the participants to be consumed by the monkeys from their cultivated land or gardens.

Local name in Scientific name Monkey species reported to Respondents Spanish consume or damage (%) (No. respondents) Abiu Pouteria caimito Cebus (1), Alouatta(2), Ateles(1) 17 banano morado Musa spp. var. Cebus (1) 4 Biribá Rollinia deliciosa Ateles (3) 13 Cacao Theobroma cacao Cebus (4), Alouatta (1) 21 Caimito Chrysophyllum Cebus (1), Alouatta (6), Ateles (1) 33 cainito Guayaba Psidium guajava Cebus (3), Alouatta (4), Ateles (2) 38 guava cuchilla unknown Cebus (1) 4 Maíz Zea mays Cebus (1), Ateles (1) 8 mamón chino Nephelium Cebus (2), Alouatta (5), Ateles (4) 46 lappaceum manzana de agua Syzygium Cebus (3), Alouatta (11), Ateles (4) 75 malaccensis Maracuyá Passiflora edulis var. Cebus (1) 4 flavicarpa Nancite Byrsonima crassifolia Alouatta (2) 8 Pejibaye Bactris gasipaes Alouatta (1) 4 Pipa Cocos nucifera Cebus (1), Alouatta (damage to 8 leaves) Plátano Musa balbisiana Cebus (1) 4 yuplón/jobo Spondias dulcis Alouatta (1), Ateles (1) 8 zapote mechudo Licania platypus Cebus (1), Alouatta (3), Ateles (2) 25

35 by participants. Not all monkey species were reported to eat each item. Overall, Cebus was reported to consume a total of 13 different species, while Alouatta was reported to consume 11 different species and Ateles, nine different species. The top three food species consumed by

Cebus as reported by participants were cacao (Theobroma cacao), guayaba (Psidium guajava), and the water apple (Syzygium malaccensis). Alouatta was more frequently reported to consume the water apple, caimito (Chrysophyllum cainito), and rambután (Nephelium lappaceum). Ateles was more commonly reported to be eating rambután, water apple, and biribá (Rollinia deliciosa).

The water apple or rambutan was in the top three food items consumed by the three monkey species as reported by participants. Similarly, rambutan was in the top three more commonly reported food items to be consumed by Ateles and Alouatta.

Perceptions and attitudes toward monkeys

A total of 30 different adjectives used during interviews to describe monkeys were identified and classified as positive, neutral or negative (See appendices D and E). Of those 30 adjectives across 67 responses (Table 10), Alouatta was described using 16 adjectives, Ateles was described using 17 of those adjectives, while Cebus was described only using nine adjectives. An association was found between monkey species and perception of monkeys (X2[4,

N=67] =15.09, p=0.0045). Alouatta is more likely to be described negatively, Ateles more positively, and Cebus was more likely to be described using neutral adjectives (Figure 5).

Interestingly, I found no association between attitude and being reported as a crop raider (X2[2,

N=67] =3.88, p=0.1435).

An association was found between location of residence and the use of positive, neutral, or negative adjectives to describe monkeys, (X2[4, N=67] =24.6, p < 0.0001). Respondents living

Table 10 The number and percent of positive, neutral, and negative responses when describing the Gandoca monkeys.

Adjective No. of Percent of Alouatta Ateles Cebus Classification responses responses Positive 22 33% 8 11 3 Neutral 19 28% 3 5 11 Negative 26 39% 13 6 7 Total 67 100% 24 22 21

80%

60%

40%

20%

0% Alouatta Ateles Cebus

Positive Neutral Negative

Figure 5 The percent of responses that were positive, neutral, or negative for each monkey species. Each species had a different total amount of describing responses: Alouatta (n=24), Ateles (n=22), Cebus (n=21).

near the beach were more likely to describe monkeys positively, while people living in the

Pueblo area were more likely to describe monkeys negatively, and people living in Bonifé were more likely to describe monkeys using neutral adjectives. Moreover, women were more likely to describe monkeys negatively, while men were more likely to describe monkeys with more

37 neutral or positive adjectives, (X2[2, N=67] =15.08, p=0.0005). This could go in hand with the fact that women were more likely to report crop raiding to be a problem (X2[1, N=24] =8.00, p=0.0047)

When participants were asked if they had preference of one monkey species over the other, most responded to have no preference (44%) for any of the species (Figure 6). However, four of the participants who had no preference also explained that they do not like monkeys, they do not pay much attention or that they like them all equally.

Ateles 33%

No preference 44%

Cebus 19% Alouatta 4%

Figure 6 Respondents' (N=24) preference for the different monkey species.

Management practices

Participants (N=24) were asked about any mitigation methods and/or practices for preventing crop raiding that they use in general with the problematic animals they reported.

Respondents reported a total of 12 different methods (across 45 responses) for preventing different animals from eating or damaging their crops and/or gardens (Table 11). However, only seven of those methods targeted monkeys. The perceived success for each method was calculated

38 according to the percentage of participants who stated the practice to be successful. If less than

25% of the respondents reported the methods to be successful, then it was classified as a method

Table 11 The number and percent of respondents for each mitigation methods against problem animals and crop/garden raiders and their level perceived success.

Measures taken against Target animal No of Percent Perceived problematic wild respondents respondents Success animals (%) Pesticide insects 1 4 High Planting fruit trees in monkeys, birds 4 17 Low forest Buffer strips of fruit trees monkeys 2 8 Moderate Killing birds, rodents, 3 13 High monkeys Guard dogs monkeys, 9 38 Moderate predators Cutting connective trees monkeys 8 33 Low Scarecrows birds 2 8 Moderate Scare them away monkeys, birds, 5 21 Low small mammals Making noise with birds 2 8 High aluminum cans Mirrors in the trees birds 2 8 Moderate Arrows squirrels 1 4 High Pyrotechnic monkeys, birds 1 4 High None N/A 5 21 -

of low perceived success. If more than 25%, but less than 75% of the respondents claimed the method to be successful, then it was classified as of moderate perceived success. Finally, if more than 75% of respondents reported the method to be successful, then it was classified as of high perceived success.

The majority of people reported to have dogs to guard their chickens and to scare away monkeys, with a moderate perceived success. Another common practice to help prevent monkeys

from entering their land is to cut trees that may connect

the forest to their gardens and/or crop land (33%). Other

practices that are specific against monkeys are

strategically planting food lures in the forest (17%) and as

buffers around crops (8.3%) like cacao, coconut, and water

apple (when produced for sale). Killing was directly

reported for animals such as the tepezcuintle, the gavilán,

and for monkeys (indirectly). For example, one participant

mentioned that once you kill one spider monkey you will

not see them come back for years because they remember.

Dogs were also reported to kill monkeys if they come

down from the trees. Some participants (21%) claimed not Figure 7 Scare crow installed in the canopy of a rambutan tree to scare to use any methods against crop raiding monkeys because away raiders, only reported to work for oropendolas. it is not necessary or because it is not successful.

The most common practices against monkeys were only perceived to be low and moderately successful (Figure 8). Participants explained that planting fruit trees for monkeys in the forest helps, but it does not prevent them from entering their land in search for more.

Similarly, creating buffer strips of other fruits to protect their crops was moderately perceived as successful. Only one respondent stated that strategically planting other fruit trees has helped them protect their crops. Although cutting connective trees is a popular practice, it is described as unsuccessful for all monkey species. Many of the respondents explained that if monkeys want to get to a food source they will come down from the canopy and walk to the source. In this case, if dogs are not on a leash, they might attack and kill the monkeys.

100%

80%

60%

B 40% A

A 20% A B

0% Planting fruits Buffer fruit Guard dogs Cutting Scare them in the forest trees connective trees away

Figure 8 The methods used by respondents to prevent or deter monkeys from entering their land and/or raiding their crops and gardens and the percent of respondents (N=24) who used each method (A=low perceived success; B=moderate perceived success).

Importance of conservation

Almost all participants, with the exception of two, indicated that wildlife conservation is important. Respondents were asked the reason behind their answer and responses (n=33 responses) were classified into six groups (Table 12). Nearly half of respondents explained that wildlife conservation was important because one must help maintain the balance of nature and every animal has an ecological role (44%). Approximately a quarter of the participants (26%) explained that wildlife conservation was important for our own benefit and entertainment, while

22% stated that it was important to preserve so that our future generations can enjoy it as well.

Another 22% of respondents said that the conservation of wildlife is important for the purpose of ecotourism. Lastly, 17% said it was important to take care of God’s creation and 13% said because we should live in harmony with nature.

Table 12 The reasons for wildlife conservation as stated by respondents (N=24; n=33 total responses).

Reasons for wildlife No. Percent No. respondents who conservation respondents respondents benefit from tourism Tourism 5 21.7 3 Ecological value 10 43.5 4 For future generations 5 21.7 1 Utilitarian value 6 26.1 2 Religious 4 17.4 0 To live in harmony with nature 3 13.0 2

The reasons given by the respondents (N=24, 33 responses) were then assigned three different values (Pearson 2016). The three different values were: intrinsic, utilitarian, and non- use (Figure 9). More than half of the reasons given for wildlife conservation were of utilitarian

value (64%), such as for tourism, Intrinsic. 18% fulfilling an ecological role, or for

human benefit. The rest were of

intrinsic value (18%) and non-use Non-use. 18% value, such as for the use of future Utilitarian. 64% generations or simply conserving

for the goal of living in harmony Figure 9 Percent of responses (n=33) for the importance of conservation that had a utilitarian, intrinsic, and non-use value. with nature.

Observations of feeding instances and camera traps

As part of the study design, vigils were to be conducted to make observations of monkey

crop raiding behaviors and compare actual crop raiding behavior frequency and intensity.

However, crop raiding by monkeys was not as common as expected, and monkeys were not well

habituated. Nonetheless, opportunistic observations were made of Ateles and Cebus consuming

different food items.

Ateles was observed consuming the fruit from the

yolillo palm (Raphia taedigera) in the Gandoca

Lagoon (Figure 10). During this observation an adult

female and her infant were seen feeding. Images and

videos were also captured of A. geoffroyi ornatus

eating mamón chino (Figure 11), also known as

rambután, a nonnative species from Eastern Asia.

Videos showed at least three spider monkeys eating

ripe and unripe rambutan. Ateles was also observed

consuming water apple (Syzygium malaccensis) from a

garden in the Playa region. On the other hand, Cebus

was observed feeding from the red mangrove leaves Figure 10 Yolillo fruit (Raphia taedigera) half- eaten by Ateles. and the banano morado (Musa spp.). Cebus was also

observed taking a coconut from a palm tree and trying to break it open. However, once he

noticed he was being watched, the entire group fled. Alouatta was not observed eating fruits or

crops, but groups were seen near banana plantations resting (Figure 12).

Figure 11 A black-handed spider monkey (A. geoffroyi ornatus) feeding on rambutan fruits (Nephelium lappaceum) captured by a Stealth Cam camera trap. A video showed a group feeding from both ripe and unripe fruits.

Figure 12 Group of howler monkeys (Alouatta palliata) resting near a banana (Musa spp.) plantation.

CHAPTER IV

DISCUSSION

In my study of the human-primate interface in Gandoca, Costa Rica, I aimed to determine which behaviors may affect people’s perceptions and attitudes towards monkeys and whether a conflict existed between monkeys and people as reported through interviews.

Although very few people said crop-raiding was problematic, the majority indicated that monkeys do visit their land to eat from their crops and/or gardens. Participants who said it was problematic might have been experiencing more intense and frequent visits from monkeys. However, people were bothered mostly by how wasteful monkeys were when feeding rather than the feeding itself.

Monkeys are among the most problematic animals in Gandoca

According to the participants in my study, crop raiding was not a significant problem.

However, 23 animal species were reported to crop raid and/or feed from gardens frequently.

Primates and birds head this list, with Alouatta as the most frequent visitor to raid crops and/or gardens according to the participants. Squirrels, as well as Cebus, follow Alouatta in this list. Even though Ateles was not reported to raid gardens as frequently as other species, it was more commonly reported to be the most problematic raider amongst the visitors.

Alouatta are known for their ability to persist in spite of extreme anthropogenic disturbance to their habitats and thrive in matrix habitats (Garber et al. 1999; Arroyo-

Rodríguez and Dias 2010). I predicted Alouatta to be described as the most problematic species among the three monkeys in Gandoca, because of their characteristic persistence through matrix habitats and their more conspicuous behaviors/visible presence. I also

46 expected Cebus to be a greater problem for locals and to be seen more negatively, but capuchins in Gandoca are shy around people and harder to see. Although they still raid gardens and crops, they were not reported to do so as much as Alouatta. In comparison to

Alouatta, Cebus and Ateles move more quickly and are more agile, so can be harder to see if they were raiding. As I predicted, Ateles is the least frequent raider of the three monkeys and the more positively perceived. Although they heavily rely on fruits, so you would expect them to come more for the crops and cultivars in the gardens, they are not well habituated, often running away from human presence. Spider monkeys also rely more on undisturbed habitat, spending more time in the forest canopy (Aureli and Schaffner 2010). However, people still reported all three species to move terrestrially to get to their land after cutting connective vegetation.

Previous studies have reported all three genera to crop raid (Alouatta: McKinney

2010; Ateles: Waters and Ulloa, 2007; Cebus: Galetti and Pedroni 1994, Baker and Schutt

2005, De Oliveira and Fialho 2007). However, the most common of the three and often described as a destructive forager is Cebus (Terborgh, 1983). Alouatta has been previously described as wasteful while feeding, as well as Cebus and Ateles (Howe 1980; Estrada and

Coates-Estrada 1984)

Perceived differences between monkeys in Gandoca

Most of the people interviewed in my study stated they had observed some behavioral differences between the sympatric alloprimates. Overall, monkeys were described as harmful animals to crops and/or gardens and to be wasteful when feeding. However, Alouatta was perceived to be more so than Ateles and Cebus. Although Alouatta was the most frequently

47 reported raider, it was not necessarily the most problematic for the people who reported them. Participants described Alouatta to be calm, a slow traveler, friendly with people, and tame, but also lazy, and very wasteful and harmful when feeding from their cultivars. Many participants explained that Alouatta stays for a longer period of time on their land when passing by and that they also leave behind bad odors. In contrast, Ateles was described as agile, pretty, fun to watch, and while it is often curious of people it is still very shy and afraid of people. Although still described as wasteful, participants explained that Ateles tends to visit their land for shorter periods of time, taking only what they need and returning to the safety of the forest to eat it and often returning every day for their favorite ripe fruit available. On the other hand, descriptions of Cebus feeding habits were not as detailed.

Participants mostly described Cebus as shy with people, but a bandit and a wasteful raider.

Capuchins have been previously described as “destructive” foragers (Terborgh, 1983) due to the techniques they use while searching for and trying different foods. They have also been described as the most problematic crop pest in Costa Rica (Baker and Schutt 2005).

However, this does not seem to be the case in Gandoca.

A possible explanation for Alouatta being the most common raider, as reported by participants, is that it is more widely distributed through Gandoca, and it is easier to observe given that it moves slowly and stays for a longer period of time in an area. They are also less fearful of humans. Conversely, Cebus is harder to observe given that it moves quickly through the trees and avoids people at all costs.

“Monkeys love the water apple”

Respondents listed a total of 17 different food items, including crops, and native and

nonnative cultivars. Cebus was reported to feed from 13 different items, mostly of cacao, guayaba, and water apple (also known as malay apple). Alouatta and Ateles were reported to feed from fewer items than Cebus. Alouatta was reported to feed from 11 different items of which were mostly water apple, caimito, and rambután. Ateles was reported to feed from nine different items, which were mostly rambután, water apple, and biribá. Although some of these are not crops, some respondents explained that when it is possible they do sell these fruits. Interestingly, some participants expressed that they prefer monkeys and other animals to eat the fruits that people are not consuming as much, because otherwise those would go to waste. For example, Rollinia deliciosa was described as a delicious and nutritious fruit, but locals do not consume it as much, and as a result it may create a “mess of rotten fruit” in their property. Again, this may prove that “gandoqueños” do not necessarily care if monkeys eat from their land, but what and how they are eating it. It is important to further study the feeding habits and confirm the consumption of these items by the sympatric primates of

Gandoca. Determining how significant these food items are in their diet and ecology is crucial for the adequate management of raiding species (Hill et al. 2002).

A number of the participants, mostly people who have been living for more than 30 years in Gandoca, described how the diet of the howler monkey has changed. According to them, Alouatta used to have a more fruit-based diet and now they are eating mostly leaves due to forest loss. The also explained that given that they are the slowest and “not so clever” of the three species, they have not been able to compete for fruits with Ateles and Cebus because they are agile and they can overpower them. The fact that people are giving such an explanation demonstrates how observant they are of the animals or how close they are to

49 monkeys. As previously mentioned, Alouatta is known for its flexible behavioral and ecological traits, making it easier to adapt to anthropogenic habitats (McKinney et al. 2015).

Perceptions and attitudes about monkeys

In general, monkeys seem to be perceived as destructive and wasteful when feeding, clever, and even pretty. As expected, participants seem to have different attitudes towards each monkey species, having a more positive attitude towards Ateles, more neutral towards

Cebus and more negative towards Alouatta.

Previous studies have found that communities may feel differently towards different animals, and these attitudes can be influenced by behaviors and physical traits displayed

(Costa et al. 2013; Sousa et al. 2014). In addition, if the animal affects the community’s livelihood, people are more likely to hold negative attitudes towards the animal and its conservation (Campbell-Smith et al. 2010). However, in this study, the participants’ attitudes towards the monkeys are not associated to the monkeys being reported as garden/crop raiders. In Gandoca, Alouatta was more often described as a destructive and damaging animal, while Ateles was more often described as pretty and Cebus as shy of people. A great variety of behavioral and physical traits seem to be influencing people’s perceptions and attitudes about primates, such as their feeding habits, communication, interactions with people, smells, and sounds, among other traits.

People who claimed monkeys were problematic and crop/garden raiders still described them positively. Many respondents expressed that it was not the animals’ fault; monkeys are running out of food in the forest because their homes have been deforested, and they have to eat what they find. A number of participants said that “animals have the right to

50 feed from the forests, it is their land as well”. Additionally, many of the participants expressed that they do not necessarily care if monkeys feed from their crops or gardens, but they are bothered by how much they “waste” and “damage” when they feed. They explained that they sometimes bite a fruit and throw it away without consuming the whole food item.

This perception towards “wastefulness” has been found in other studies and it was stated to affect people's attitudes (Sousa et al. 2014; C. M. Hill and Webber 2010). Sousa and colleagues (2014) found chimpanzees to be described by locals as “not bad” crop raiders.

Similar to this study, an animal can be an acceptable crop raider as long as its behavior minimizes waste and damage. Using human characteristics to describe primate behaviors is easy to do given the physical and behavioral similarities we share. Although this can help primate conservation efforts, it can cause people to have a set of expectations similar to the ones for humans and in effect, it may cause people to have a negative attitude when those expectations are not fulfilled.

Participants who live near the Playa region of Gandoca were more likely to have a positive attitude when describing monkeys. This could be due to their increased benefit from ecotourism. People who live in this region own cabinas, sodas, pulperías, and offer surf lessons and guided tours inside the wildlife refuge, therefore benefiting more directly from tourists. Tourists come to the Gandoca sector of the refuge in order to experience the uniqueness of this community living so closely with nature and wildlife. The locals living in the Playa region recognized that if monkeys and other wildlife did not exist as it does in

Gandoca their livelihoods would be greatly affected, since a great portion of their economy depends on ecotourism.

On the other hand, participants who lived in the Pueblo region had a more negative attitude and were more likely to report monkeys as raiders. The Pueblo region relies a lot more on agriculture in comparison to the Playa region, since tourists do not want to stay far from the beach. This more negative attitude could indicate that this region is likely struggling more with raiders than other regions of the community given their economy is more dependent upon subsistence agriculture and receives no direct benefit from tourism. In comparison, the Bonifé region was more likely to have a neutral attitude towards monkeys even though nearly half of the respondents residing in this region reported monkeys as raiders. In Bonifé the economy is still highly dependent upon agriculture but more for sale than for their own consumption. For example, a participant from Bonifé reported many animals to raid their garden and crops, including the three species of primates, but he said that since they produced a lot of different food items in large quantities it does not bother him because it is not affecting him significantly. However, he explained that if the conflict were to get more intense, his response would be different. Another important reason is that in

Bonifé, some of the plantations and monocultures were significantly further from the forest in comparison to the other regions. Monocultures are not common in Gandoca, at least not in large quantities, but in Bonifé it is more common to see monocultures of Cocos nucifera,

Musa spp, Theobroma cacao, and Cucurbita species.

“Wasteful” behaviors by American primates while feeding have been previously observed (Howe 1980; Estrada and Coates-Estrada 1984; Burton and Carroll 2005).

However, it is not yet clear why primates “waste” fruits. I thought one explanation could be tied to their vision and that dichromatic primates, those who only distinguish between greens and blues, were more likely to waste fruits while they search for ripe fruits. Male spider

52 monkeys and capuchins have a higher probability of having dichromatic vision, than females of their species (Hiramatsu et al. 2005). On the other hand, howler monkeys mostly have trichromatic vision (Jacobs et al. 1996), and they are still reported to “waste.” Another possible explanation for fruit “waste,” is that of consuming the most nutritious portion of the fruit and discarding the rest to maximize nutrient intake, sometimes called high-grading. For example, brown bears (Ursus arctos) are known to sometimes consume the most nutritious parts of their food, such as the brains and eggs of a salmon, to maximize their weight gain or to eat fast before thieves get to their food (Robbins et al. 2007; NG 2016). An alternative explanation to fruit waste is that of the flavor of the fruit. Monkeys could be trying or tasting fruits by taking a bite and discarding what they don’t like. For example, different levels of tolerance for antifeedants are might affect fruit choice and levels of consumption in primates

(Wrangham, et al. 1998). Finally, a larger scale possible explanation for fruit waste is that of seed dispersal (Howe 1980; Estrada and Coates-Estrada 1984).

In this study we found that women were more likely to have a negative attitude towards monkeys while men were more likely to have a more neutral or positive attitude.

Similarly, Priston (2005) found women to have less tolerance towards wildlife and suggested that it could be related to being unsuccessful at deterring raiding species. However, both men and women in Gandoca seem to be as unsuccessful in this task. A study conducted by

Gillingham and Lee (1999) also found women more likely to exhibit negative attitudes towards wildlife conservation potentially because of their marginalized position in community issues. In the case of Gandoca, women are involved in community issues and many are leaders who help make decisions. In another study, Ellwanger et al. (2014) found women to have a higher rate of anthropomorphic feelings toward animals. Anthropomorphic

53 descriptions were commonly used by both men and women in Gandoca, but women were more likely to do so. Perceiving monkeys as more human-like animals could lead people to have certain expectations, similar to the ones we have for humans. When monkeys do not meet those social rules, and act “lazy” and “wasteful”, it may lead people to have a negative attitude. Perhaps women having more humanistic perceptions towards animals is negatively affecting their attitudes in Gandoca.

"Spider monkeys are my favorite monkey"

Although nearly half of the respondents stated to have no preference between the three monkey species, the respondents who did express preference said to prefer Ateles.

When asked why, most of the respondents explained that Ateles is more fun to watch, prettier, and smarter. Even a participant who reported Ateles to be the most problematic raider, explained that Ateles was his favorite monkey because it is beautiful. This same participant, previous to the interview, had jokingly expressed that he was going to kill the spider monkeys raiding his garden and cook them Caribbean style. However, when privately talking to him he expressed respect to wildlife and no interest in eating monkey meat.

Some participants also expressed preference over Cebus, although less frequently than Ateles, and explained that they were pretty and smart. On the other hand, only one participant expressed that Alouatta was his favorite species and he explained that it was because this species was the one who visited him more often. At the same time, he had described Alouatta negatively, as a lazy and wasteful creature. In spite of negatively describing howler monkeys, he explained that it was important to protect God's creation and respect every creature. Religious beliefs are often considered an important factor affecting

54 people's attitudes toward conservation efforts and it may help maintain a positive attitude of conservation.

Perceiving primates as human-like creatures

A very common story told among people in Gandoca was that of white-faced capuchin monkeys being incredibly clever animals. The anecdote goes as follows:

“In Guanacaste, capuchins can be very problematic for maize growers. They go in groups to the corn fields and they work as a team. One will work as a sentinel to keep an eye out for the farmers, while the others are getting two husks of corn. They tie together the husks to free their hands, and quickly run back to the forest to eat it in safety and share with the rest of the team.”

This anecdote seems to have traveled across the country; however, some Gandoca residents claim that this does not happen in their community, but in the northwestern area of the country. Baker and Schutt (2005) stress the importance of taking folktales like this seriously.

Although it points out the intelligence of capuchins, it can be to their demise if people perceive them as animals hard to control due to this same intelligence.

On the other hand, there are certain behaviors and physical traits that can increase tolerance towards wildlife. For example, a participant pointed out that Ateles looks like it is

“blowing kisses” and it reminds him of the way humans behave to demonstrate attraction or affection, but the monkey is displaying an embrace behavior. Another example identified during interviews were maternal behaviors. Participants, without being asked, shared that they felt certain connection and pity towards these animals because “they care for their young just like we do” and some even mentioned that they eat certain fruits the same way humans do.

Importance of wildlife conservation

Nearly all participants said that conservation is indeed important, with the exception of two participants who responded that it "depends." Nearly half of the participants who responded positively to the conservation of wildlife explained that animals must be protected because they each have a purpose in nature by keeping the ecosystem in balance. Another common response was based in animals having use for humans or an indirect benefit to us, for example entertainment, food, maintaining mosquito populations under control, among other things. Participants also expressed that if wildlife is conserved future generations can enjoy and benefit from them, as well as tourists, which bring money to the community. The least common responses when participants were asked why conservation is important were based on religious beliefs, and because humans must simply live in harmony with nature and respect it. Although participants were not asked directly about their religious beliefs,

Christianity is the main religion in Gandoca. However, witchcraft was also mentioned to be practiced by some, but nobody claimed to do so since it appears to be condemned and frowned upon by many in Gandoca.

Moreover, the most common reasons given for the importance of wildlife conservation were of utilitarian value, followed by intrinsic and non-use value, as defined by

Pearson (2016). Expressions such as “what is a forest without animals?” or “a forest with no animals is like a garden with no flowers” were not uncommon. One must keep in mind the history behind the origins of the Refugio Nacional Mixto de Vida Silvestre Gandoca-

Manzanillo, Jairo Mora Sandoval. The wildlife refuge was first established through the initiative of the community to protect their land and their natural resources against foreign exploitation. Although people were allowed to keep their land through this agreement, they

are not allowed to continue expanding their land through deforestation. However, many

locals still practice ‘slash and burn’, the clearing of primary and secondary forest for new

plantations (Figure 13), but they carefully choose an area that will not be easily found by the

government authorities. While most

of the respondents claimed that

wildlife conservation was important,

two participants explained that it

depends on the situation. When asked

for a further explanation, both

participants expressed that

conservation can be practiced as long

as their livelihoods or economy are

not being affected by the animal’s

actions. Moreover, they explained

that first they will think about

improving their business or

economy, by expanding their

cultivated land for example, and then

think about conservation. Both of

these participants had claimed to be

Figure 13 The clearing of forests near their land is responsible of the death of some common to start new plantations as needed, such as for banana or plantain. animal (i.e., paca, bird of prey and

boa) in order to protect their crops and domestic animals. Although many in Gandoca seem

to support wildlife conservation, livelihoods may take precedence in cases of intense conflict.

In a similar study, conducted in Karnataka, India, societal expectations, such as following the

laws or protecting wildlife and preserving nature, were ignored once people were being

affected by wild animals’ actions (Chakravarthy and Thyagaraj 2005).

During an informal

conversation, a local woman

shared her views on

conservation and her

perception of some

environmental activists. She

stated that one thing is to take

care of nature, but some

conservationists are extremists

and fanatics. Further, she said

that “one must not be too much

of an environmentalist,” and

she continued to describe them

as people who do not want to

leave nature. She believes that

people should be progressive

and not conform to the jungle.

Figure 14 According to locals, several years ago, people from the Although this same woman Netherlands established a program where they gave monetary compensation to locals as incentive to keep their land untouched complained and reported to and use that simply to be preserved. Some people are still under that program.

58 experience intense garden and crop raiding from a variety of animals, she explained that humans are the ones to blame because we have destroyed their forests and monkeys have no choice than to feed from our cultivars. She was not the only person to explain that monkeys are only trying to get the food they are not finding in the forests and thus we have to share the right to feed from the land with other animals.

Perceived relationship of los gandoqueños with wildlife

A topic that came up in several interviews was that of hunters, known in Gandoca as

"monteadores." Hunting was recently banned in Costa Rica, only allowing hunting for personal consumption in limited circumstances, research or wildlife population control.6 In

Gandoca, many stated hunting to be an issue because people from outside the refuge were entering to hunt for different animals, including indigenous people from Panamá or people from Manzanillo, the community adjacent to Gandoca. Nonetheless, people from Gandoca also hunt for their own consumption and for sale, but it is a topic that stays quiet given that it is a wildlife refuge and it is frowned upon by other members of the community. Moreover, it seems to be less of a problem if a Gandoca resident hunts as opposed to a visitor from a nearby community.

Management practices

Nearly all participants claimed to use one or more methods to deter animals from raiding their crops and gardens. From a total of 12 different methods to discourage raiders, seven targeted primates. Having guard dogs was the most common method used by participants to prevent crop and garden raiding and the predation of farm animals. Although

6 http://www.ticotimes.net/2013/01/31/president-chinchilla-signs-ban-on-hunting-for-sport

59 having guard dogs had only moderate success, it was more successful if the dogs were loose instead of tied up. Respondents explained that if the dogs are always tied up in the same place, monkeys may learn that dogs cannot really do anything to them except for barking. In this case, monkeys would still enter gardens or cultivation using trees as corridors or terrestrially if necessary. People who claimed dogs to be successful at deterring monkeys had dogs who were loose within their property. Two respondents revealed that their dogs had killed howler monkeys that entered their property by land. Those participants felt sorry for the monkeys but still encouraged this behavior. One of the two respondents continued sharing and explained that since she would not want the monkey meat to go to waste, she would roast or grill the monkey carcass and feed it to the dogs.

Further, cutting trees or vegetation that may serve as corridors for monkeys into their land and plantations was another common method to prevent these arboreal raiders. Although common, it seemed to have low perceived success. Respondents claimed that often, all three alloprimates would enter their property moving terrestrially if they really wanted to feed from a fruit tree. Further study into why monkeys in Gandoca do not respond as expected to this method would be important to improve mitigation methods. For example, what influences a monkey’s “decision” to walk across open land to reach a certain fruit and expose themselves to predation and landowners? Could it be for some fruit rich in nutrients or more common during periods of scarcity? What makes this strategy worth the risks? Another method to prevent monkeys from raiding was to plant fruit trees in the forest, such as

Rollinia deliciosa, Nephelium lappaceum, and Syzygium malaccense. Syzygium is said to be used by coffee growers in other areas of Costa Rica to divert birds from their precious crop.7

7 en.academia.ru/dic.nsf/enwiki/11682279

Respondents mentioned that by providing them with some of their favorite fruits in the forests, they expected monkeys not to have the need to enter their land. This method had a low perceived success; however, participants who claimed to have done this reported that they would still do it just to “provide food for the monkeys”. One respondent said that her dad taught her to plant for the animals as well, because “we share the land with wildlife and we are responsible of helping them”. This may reflect the importance of educating parents and children on the importance of wildlife, conservation, and the intrinsic and utilitarian value of nature.

As previously mentioned, one of the most common explanations given by participants for garden and crop raiding by alloprimates was the lack of resources in the forest. A study conducted by Seiler and Robbins (2015) investigated the effects of food availability outside and inside of the Bwindi Impenetrable National Park in Uganda on the occurrence of gorilla ranging outside the park boundaries and raiding crops. They found that the availability of palatable crops, pines and tea plantations was what influenced gorillas to crop raid and not the lack of food in the national park forest. On the other hand, chimpanzees are known to increase crop raiding during fruit scarcity (Naughton-Treves et al. 1998; Hockings,

Anderson, and Matsuzawa 2009; McLennan 2013). Similarly, I hypothesize that Alouatta, often compared to Gorilla in behavior and ecology (Di Fiore et al., 2011), is not necessarily driven by food scarcity to crop raid, but opportunity. On the other hand, I hypothesize that

Ateles, a less common raider in Gandoca, to be more likely to raid gardens and crops in times of scarcity, as they tend to continue to include fruit in the diet during periods of fruit scarcity by ranging further, rather than falling back on other food types (Klein and Klein 1977;

Milton 1981).

Another method mentioned by participants was planting buffer crops or fruit trees that may decoy monkeys from entering further into a plantation so that they are satiated before continuing into a cultivation or garden. Respondents who use this method explained that they plant trees of some of their favorite fruits, such as Rollinia deliciosa, Nephelium lappaceum, Syzygium malaccensis, surrounding crops like Theobroma cacao, Musa balbisiana, Syzygium malaccensis and Cocos nucifera. One respondent who claimed to have used this method, said that raiding by Cebus and Alouatta significantly decreased after implementing the buffer strips as food lures, in addition to planting more fruit trees inside the adjacent forest. In this case, crop raiding decreased but they still enter her plantation when there are fruits. On the other hand, the other participant who claimed to have planted fruit trees as buffer, said that nothing seems to work for deterring monkeys from raiding his garden because “they will always find a way to get to the fruits they want”. In this case,

Ateles was the most problematic raider to enter his garden and eat Rollinia deliciosa and

Nephelium lappaceum. However, he claimed that he does not mind that monkeys eat R. deliciosa because he does not consume that fruit, but he does not like monkeys consuming N. lappaceum because he does consume that fruit and not only monkeys are feeding from it, but also oropendolas and birds of the Psittacidae family.

Buffer crops were recommended in Bwindi Impenetrable National Park to deter gorillas from raiding important crops (Seiler and Robbins 2016). In this site, the buffer crop was unpalatable for gorillas, planted over large areas, and preferably economically valuable for farmers. In addition, it was recommended to be cleaned of other vegetation that may attract gorillas. In Gandoca, crops that are palatable for monkeys are used as buffers instead.

In the Refugio de Vida Silvestre Curú in Costa Rica, a similar method was used and was

62 called “food lures” (Baker and Schutt 2005). Food lures, such as a plantain species (Musa spp. var. filipita) are meant to draw wildlife away from cash crops like melon. The species of plantain used in Curú is not as valuable for locals as it used to be, but the fruits provide important nutrients, the plants holds water, provides shade, nectar, and it is home to a variety of insects, making it the perfect lure for Cebus in this site. Although, this Musa variety has been proved to be successful for Cebus, each population should be examined individually for the best buffer crop and strategy that can mitigate crop raiding by primates.

It is important to keep in mind that (almost) no method will completely eliminate crop raiding, but it will likely mitigate some of its effects. Deterring primates from raiding crops and gardens has been challenging. Non-human primates, our closest living relatives, have physical and behavioral traits that make them difficult to manage. Developing strategies that not only treat the symptom but target the underlying causes for crop raiding is vital for successful management (Jackson et al. 2008).

Recommendations

Before implementing any preventive methods against crop raiding, it is essential to understand the context of crop raiding (Sitati et al. 2005); the frequency and extent of the raiding events, crops grown on farms, and the behavioral ecology of the species that are raiding (Hill et al. 2002). Also, it is important to communicate with the residents and to assess their perceptions of wildlife, conservation, and of crop raiding as a conflict

(Gillingham and Lee 1999). Finally, consulting with the local community to make sure their needs and goals are taken into account while developing any management plans (Webber et al. 2007).

In Gandoca, not all people consider crop raiding to be a problem. Nonetheless, many expressed interest in learning about new methods to deter or prevent raiding primates from entering their lands and eating from their plantations and gardens. And as plantations continue to replace forests in the Americas, we must not oversee crop and garden raiding and studying human-wildlife interactions. Next, I will present recommendations based on findings from this and previous studies. I recommend the use of more than one of these methods as necessary and approach these recommendations as cost-benefit scenarios. For example, cutting connective trees can be a first step to eliminate the canopy bridge between the forest and the farm, but complementing with hedges or living fences that deter or repel monkeys from entering. Also, identifying the points of entrance for monkeys into the farmer’s land, can help determine what measures need to be taken. A warning system could be developed using ropes and bells near plantations to alert the farmer of the presence of raiders and the approximate location. Although some of these may be relatively inexpensive, they may be costly if farmers are required to constantly spend too much time with little benefit.

Hedges and barriers

Hedges or living fences are composed of strips of vegetation that form a barrier and keep crop raiding animals out. A study conducted by Hill and Wallace (2012) tested the effectiveness of 10 deterrents in four different Ugandan villages against crop raiding primates. They found several barrier systems to be effective and to reduce the frequency of crop raiding events, such as four-strand barbed wire fences and a combination of ocimum

(Ocimum kailimandscharicum) hedge and mesh fence. Ocimum is a perennial, aromatic

64 shrub native to some African and Asian countries,8 growing between 0.8-1.2 meters. This basil species is believed to be an effective deterrent because of its aromatic smell, and the high amount of bees it attracts, in addition to the way it forms a dense and interwoven barrier. Cebus apella has been previously observed to respond aversively to strong and unusual odors (Ueno 1994). Therefore, incorporating similar aromatic herbs found locally into barriers or living fences could be a potential primate deterrent in Gandoca, or at least to be tested. Additionally, some of these plants could have an economic potential or simply a domestic use.

Another hedging material proven effective by Hill and Wallace (2012) was jatropha

(Jatropha curcas). Jatropha is a drought resistant shrub native to America and sometimes used as living fence to manage livestock (Makkar et al. 1997). It may reach a height of 8-10 meters in favorable conditions, but it usually attains approximately 6 meters of height9. In addition to blocking the primates’ view of crop fields, the seeds of jatropha are toxic and may add to its deterrent impact. Different parts of the jatropha plant have been reported to be used in folk medicine, cooking, poison, to mention only a few, as well as for biofuel11. Jatropha is found in Central America and it is known as nuez de Barbados, física o piñon. I recommend the plantation of these at the border of plantations or garden as a living fence or even as a buffer zone, in combination with the clearing of connective trees.

Buffer Zones

Buffer zones are the areas of land between cultivated areas and the animals’ habitat.

These are usually planted with unpalatable crops, such as tea (Rode et al. 2006), that may

8 http://apps.kew.org/wcsp/namedetail.do?name_id=136967 9 https://www.hort.purdue.edu/newcrop/duke_energy/Jatropha_curcas.html#Description

65 have economic value and discourage raiding animals from entering plantations, or even chili plants. The width of buffer zones has been suggested to be as large as 500 meters (Naughton- treves et al. 1998) and as small as 20 meters in width (Yuwono et al. 2007). The width of buffers zones depends on the animal intended to be deterred and also on how much land a farmer can dedicate to this. Because of how variable recommendations are, I suggest starting with approximately 20 meters and increasing if necessary and possible.

Repellents: spent coffee grounds

Caffeine has been tested for taste tolerance in different primates, including Ateles geoffroyi (Laska et al. 2009; O’Brien 2013), and monkeys were found to have low tolerance threshold to the taste and smell of coffee. Given that each household possibly consumes at least one cup of coffee a day in Gandoca, the spent coffee grounds could be saved and later used as a potential repellent of raiding animals. It is suggested to place these on the boundary of the farm that is located closest to the forest edge where the point of entrance is and where raiding most likely occurs (Naughton-Treves 1998; O’Brien 2013). Additionally, spent coffee grounds are known to be an effective organic fertilizer and deterrent of insects from crops (Cruz et al. 2012; Caetano et al. 2014). However, the use of spent coffee grounds does need to be tested in the field at deterring wild primates and the amount that is needed for its effectiveness.

Future Directions

Conservation efforts and wildlife management practices require community involvement for it to be successful. Therefore, my next step will be sharing the results of this study with the community leaders and MINAE officials representing Gandoca. Educational

66 workshops for the residents will be offered in order to share the different methods to deter crop raiding primates. Also, residents will have the opportunity to share openly with each other, methods that have been successful for their farms against crop raiding animals. In addition, I will aim to influence attitudes and perceptions by sharing new information about the Gandoca primates. Although many of the participants had positive and neutral perceptions and attitudes towards monkeys and their conservation, some still hold negative attitudes. Pickens (2015) indicated that one method used to change a person’s attitude and therefore their behavior, is to provide them with new information that addresses the cognitive and emotional components. And so, perhaps sharing the benefits of having monkeys around and explaining some of their behaviors could be a good start.

Further research is needed to better understand the complete context of crop and garden raiding in Gandoca. Following up with the Gandoca residents, studying the primate groups in Gandoca, and study the farms and gardens being raided to test potential mitigation methods.

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APPENDIX A

INSTITUTIONAL REVIEW BOARD APPROVAL LETTER

APPENDIX B

PARTICIPANTS’ SOCIO-DEMOGRAPHIC DATA

Subject Sex Age Household Household Length of Born in ID Range Size Location Residency Gandoca (yrs) (yrs) 1 Female 41-50 3 Playa 40 Yes 2 Female 31-40 4 Playa 37 Yes 3 Male 31-40 5 Playa 35 Yes 4 Female 61-70 4 Playa 38 No 5 Male 31-40 1 Playa 34 Yes 6 Male 41-50 4 Bonifé >45 Yes 7 Male 61-70 2 Bonifé 17 No 8 Male 31-40 4 Bonifé 35 Yes 9 Male 41-50 1 Bonifé 2 No 10 Male 61-70 1 Bonifé 30 No 11 Male 41-50 1 Bonifé 5 No 12 Male >71 2 Pueblo 20 No 13 Female 18-25 5 Pueblo 20 No 14 Female >71 2 Pueblo 70 Yes 15 Female 51-60 4 Pueblo 50 Yes 16 Female 18-25 3 Pueblo 19 Yes 17 Male 31-40 2 Pueblo 30 Yes 18 Female 31-40 3 Pueblo 30-35 Yes 19 Male 31-40 ND Bonifé 2 No 20 Female 61-70 5 Pueblo >60 Yes 21 Female 41-50 3 Pueblo 40 Yes 22 Female >71 3 Pueblo >70 Yes 23 Female 31-40 4 Pueblo >30 Yes 24 Male 41-50 3 Playa 11 No

APPENDIX C

PARTICIPANTS LAND INFORMATION

Subject Land under Cultivate Approximate Fruits and crops reported to be ID cultivation for sale, number of present in the participant’s land (ha) consumption, fruits/crops both cultivated 1 1 Sale 3 maracuyá, plátano, cacao 2 coco, manzana, guanábana, mango, ayote, yuca, plátano, 0.75 Both 8 banana 3 1.5 Sale 3 maracuyá, cacao, plátano 4 mamón chino, piña, yuca, pipa, naranja, manzana, maracuyá, 1 Both 8 arazá 5 cacao, manzana, mangostán, 1 Both 7 mamón chino 6 manzana, pejibaye, cacao, plaáano, ayote, yuca, mamón 40 Both >10 chino, guayaba, caña, caimito 7 plátano, naranja, tamarindo, limón dulce, limón agrio, manzana, coco, aguacate, guanábana, fruta 0.5 Both >11 de pan, piña 8 mamón chino, coco, pejibaye, cacao, guanábana, zapote criollo, zapote colombiano, limón dulce, 5.5 Both >9 naranja 9 plátano, coco, yuca, maíz, chiles, 3 Both 6 piña 10 coco, mandarina, mamón chino, 2 Both > 4 naranja 11 11 Sale 4 coco, cacao, plátano, ayote 12 maíz, cacao, mamón chino, guanábana, limón, mandarina, ND Both >7 plátano, mangostán 13 maracuyá, cacao, mangostán, mamón chino, guanábana, pipa, 1 Both >7 fruta de pan 14 cacao, guanábana, mamón chino, 1 Both >7 mangostán, caimito, abiu, maíz 15 cacao, mamón chino, biribá, zapote colombiano, naranja, carambola, guayaba, papaya, 1 Consumption 9 manzana

16 mamón chino, naranja, mandarina, 1 Consumption 5 limón, aguacate 17 mamón chino, bacurí, canela, anona, guanábana, jackfruit, ND Consumption 8 biribá, pejibaye 18 plátano, yuca, maíz, mamón 0.75 Both >7 chino, caimito, manzana 19 manzana, pipa, plátano, pejibaye, 3 Both 8 coco, limón, noni, guayaba 20 mamón, guanábana, nancite, 1.5 Consumption >5 guayaba, manzana 21 naranja, limón, mamón chino, mamón criollo, maracuyá, manzana, sincuya/soncoya, pipa, 5 Both >11 cacao, plátano 22 cacao, maíz, pipa, mamón chino, 20 Both >8 manzana, abiu, guayaba, caimito 23 0.5 Both 1 plátano 24 7 Both >5 plátano, cacao, guanábana, yuca

APPENDIX D

ADJECTIVES USED BY PARTICIPANTS TO DESCRIBE THE MONKEYS OF

GANDOCA AND THEIR CLASSIFICATION

Adjectives in local Adjectives in Classification No. of Percent of Spanish English Responses respondents (%) ágil/rápido agile/fast Neutral 2 10 alejado/arisco untamed Neutral 13 65 amigable/sociable/ friendly/social Positive 3 15 simpático bandido bandit Negative 4 20 bonito/lindisimo pretty/beautiful Positive 10 50 calmado/tranquilo calm/tranquil Positive 2 10 charlatán/travieso charlatan/ Negative 3 15 mischievous chistoso/entretenido funny/ Positive 2 10 entertaining conchú NT Negative 1 5 curioso curious Neutral 1 5 dañino/destructor damaging/ Negative 11 55 destructive feo ugly Negative 1 5 hediondo/cochino smelly/stinky Negative 3 15 hiperactivo hyperactive Neutral 1 5 inteligente intelligent Positive 2 10 juguetón playful Positive 1 5 manso tamed Neutral 2 10 ruidoso noisy Negative 2 10 sabio wise Positive 2 10 vago lazy Negative 1 5

APPENDIX E

ADJECTIVES USED TO DESCRIBE EACH SPECIES, THE CLASSIFICATION OF

ADJECTIVES AND THE NUMBER AND PERCENT OF RESPONDENTS WHO

USED EACH ADJECTIVE

Alouatta Adjective No. Percent Classification respondents respondents tranquilo/calmado Positive 2 8 ruidoso Negative 2 8 bonito Positive 2 8 hediondo/cochino Negative 3 13 manso Neutral 2 8 vago Negative 1 4 sabio Positive 1 4 dañino/destructor Negative 5 21 inteligente Positive 1 4 arisco Neutral 1 4 feo Negative 1 4 amigable/sociable Positive 2 8 bandido Negative 1 4 Ateles Adjective No. Percent Classification respondents respondents bandido Negative 1 5 bonito/lindísimo Positive 5 23 simpático Positive 1 5 chistoso/entretenido Positive 2 9 charlatán/travieso Negative 2 9 juguetón Positive 1 5 arisco Neutral 3 14 sabio Positive 1 5 inteligente Positive 1 5 conchú Negative 1 5 destructor/dañino Negative 2 9 ágil/rápido Neutral 2 9 Cebus Adjective No. Percent Classification respondents respondents bandido Negative 2 10 bonito Positive 3 14

82 alejado/arisco Neutral 9 43 curioso Neutral 1 5 dañino/destructor Negative 4 19 travieso Negative 1 5 hiperactivo Neutral 1 5