Are Jaguars Paying the Price of Our Thirst for Palm Oil?

Home , Neotropical otter, Northern screamer, Procyon (genus)

A study on the effect of oil palm cultivation and on the attitudes towards jaguars in the Middle Magdalena region, Colombia.

By Valeria Boron

Supervised by: Dr. Marcus Rowcliffe & Dr. Esteban Payan

September 2012

A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science and the Diploma of Imperial College London.

I, Valeria Boron, confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis.

CONTENTS

List of Acronyms……………………………………………………………………… 5

Figures and Tables…………………………………………………………………… 6

Abstract……………………………………………………………………………….. 8

Awknowledments……………………………………………………………………. 10

1. Introduction………………………………………………………………………. 12 1.1 Problem statement…………………………………………………………. 12 1.1.1 The Middle Magdalena Region………………………………………… 13 1.2 Project significance………………………………………………………… 14 1.3 Project objectives…………………………………………………………... 15

2. Background………………………………………………………………………. 17 2.1 The jaguar………………………………………………………………….. 17 2.2 Why conserve jaguars? …………………………………………………….. 18 2.3 Measuring habitat use……………………………………………………… 19 2.3.1 Jaguar habitat use………………………………………………………. 19 2.3.2 Habitat use by other species……………………………………………. 20 2.4 Attitudes towards jaguars…………………………………………………... 21 2.5 Palm oil…………………………………………………………………….. 22

3. Methods…………………………………………………………………………... 26 3.1 Conceptual framework……………………………………………………... 26 3.2 Study area………………………………………………………………….. 27 3.3 Camera trapping…………………………………………………………… 28 3.4 Habitat characteristics recording…………………………………………… 28 3.5 Interviews………………………………………………………………….. 29 3.6 Analyses….………………………………………………………………… 30 3.6.1 Camera trap and environmental data………………………………….... 30 3.6.2 Interviews data…………………………………………………………. 32

4. Results……………………………………………………………………………. 33 4.1 Camera trapping…………………………………………………………… 33 4.1.1 All species……………………………………………………………... 33 4.1.2 Species of focus……………………………………………………….. 37 4.2 Occupancy and habitat use analyses for the jaguar and the agouti………….. 41 4.3 Attitudes towards jaguars.………………………………………………….. 43

5. Discussion………………………………………………………………………... 47 5.1 Camera trapping and impact of oil palm cultivation………………………... 47 5.1.1 Species and habitat types……………………………………………… 49 5.2 Habitat use and occupancy analyses………………………………………... 51 5.3 People attitudes towards jaguars…………………………………………… 52

5.4 Evidence from results in relation to project objectives and research questions………………………………………………………………….... 54 5.5 Conclusion……………………………………………………………….… 55

References……………………………………………………………………………. 57

Appendix I……………………………………………………………………………. 66 I.I Jaguar habitat use…………………………………………………………...... 66 I.II Different names for palm oil and its derivatives…………………………….... 67 I.III The oil palm production chain: stages and products………………………...… 67

Appendix II…………………………………………………………………………... 68 II.I Characterization of the oil palm plantation concession “El Hato”………….... 68 II.II Interviews script…………………………………………………………...... 69

Appendix III………………………………………………………………………….. 72 III.I Photo-inventory of mammal and bird species recorded in the study area…….. 72

LIST OF ACRONYMS

JCU(s) Jaguar Conservation Unit(s)

IUCN International Union for Conservation of Nature

RAI(s) Relative Abundance Index (Indices)

CT(s) Camera Trap(s)

RSPO Roundtable on Sustainable Palm Oil

PA(s) Ptotected area(s)

FIGURES

Fig. 1.1 Map of Panthera’s Jaguar Corridor Initiative. Corridors of concern are corridors less than 10km in width at any point along their length. The JCUs mapped were defined in 2006. Source: Panthera (2012)..………………………………………………….…...... 12

Fig. 1.2 JCUs (green) and modeled corridors between them (red), in Colombia. The country JCUs were redefined and updated from Fig.1.1. The important part of the Middle Magdalena region for jaguar connectivity is circled. The corridor directly linking San Lucas with the Orinoquia population (most likely not in use by jaguars) is crossed. Modified from Panthera Colombia (2011)....………………………………………………….…...... 13

Fig. 2.1 Jaguar mother and two cubs photo-captured in the study area…………………...... 17

Fig. 3.1 Diagram representing the conceptual framework of the project...... 23

Fig. 2.3 From left to right: departments of Colombia with oil palm cultivation (green), an oil palm plantation, and oil palm fruit...... 26

Fig. 3.2 Map of the study area, by Carolina Soto...... 27

Fig. 3.3 Vegetation types and project activities. (a) Edge of secondary forest in El Hato, (b) oil palm cultivated area in El Hato, (c) retrieving camera trap data in El Hato, (d) conducting interviews in the community Cerrito. Photos a-b by V. Boron, c-d by C.M.Wagner……..…..28

Fig 4.1 Species accumulation curves. (a) All species; (b) mammals and birds considered separately...... 35

Fig. 4.2 Species of focus most frequently photo-captured. From left to right: Central American agouti (Dasyprocta punctuata), jaguar (Panthera onca), and crab-eating raccoon (Procyon cancrivorous)...... 37

Fig. 4.3 (a) The four felids photo-captured in the study area. From left to right: jaguar, puma, ocelot, and jaguarundi. (b) Locations of felid detection. Highlighted cells indicate detection of felids at camera stations (1-23)...... 38

Fig. 4.4 RAIs for the Species of Focus, calculated directly from the number of independent capture events and the number of trap nights for each palm edge habitat, and expressed as records per 100 trap nights...... 39

Fig.4.5 Cumulative RAIs for the Species of Focus and Felidae and prey species RAIs for the Species of Focus, calculated directly from the number of independent capture events and the number of trap nights for each habitat type, and expressed as records per 100 trap night...…40

Fig. 4.6 Daily activity patterns of felids and most common prey species (N=number of independent capture events (y axes) per time interval, expressed as % of the total number.....41

Fig. 4.7 Communities’ preference (“Yes” or “No”) regarding the sharing of lands with jaguars, expressed as percentages of “Yes” and “No” per each community………….…..…43

Fig. 4.8 Communities’ opinion on whether the jaguar is a problematic animal (“Yes”, “No” and “Could be”), expressed per each community as percentages of respondents giving each answer...... 44

Fig. 4.9 Reasons for considering the jaguar a problematic animal, expressed as percentages of respondents giving each reason...... 44

Fig. 4.10 Genders’ opinion on whether jaguar can attack people without being provocked (“Yes”, “No” and “Maybe”), expressed per each gender as percentages of respondents giving each answer...... 45

Fig. 4.11 Reasons for conserving jaguars, expressed as percentages of respondents giving each reason (this was an open question, however respondents only gave one reason)……………45

Fig. 4.12 Genders’ opinion on whether jaguar have increased (“More”) or decreased (“Less”) in number, expressed per each gender as percentages of respondents giving each answer…...46

TABLES

Table 4.1 List of species recorded by camera traps in study area and their IUCN and Regional Red List categories...... 33

Table 4.2 Number of independent photos and Relative Abundance Indeces (RAIs) per 100 trap nights for mammals and birds detected. Highlighted rows contain “Species of Focus”. Bold species name indicate species are listed in IUCN or Regional Red Lists...... 35

Table 4.3 Occupancy models for the jaguar (J1-J17) and the central american agouti (A1-A8). Covariates “vehicles”, “people”, “livestock”, and ”prey” were included as RAIs. “PR” stand for “Palm/Riparian” habitat, hence for vicinity of water...... 41

Table 5.1 Summary of project’s findings in relation to initial objectives and research questions...... 54

Word Count: 12,901

Abstract

Jaguars are declining and threatened by habitat loss, while oil palm plantations are expanding in Colombia, and their impact has not yet been studied on jaguars or on other Colombian fauna. Hence, camera traps were set in an oil palm plantation concession located in the Middle Magdalena region, regarded crucial for jaguar connectivity in the country. Cameras were set across different habitat types, to assess what species were present and with what Relative Abundance Indices (RAIs). Particular attention was given to jaguars, other felids species, their prey and any species listed in Regional or IUCN Red Lists, all species of focus. Further habitat use analyses were performed for the jaguar and the Central American agouti, through occupancy models with covariates. In addition, because the region is a human dominated landscape and jaguars are considered a species with a high conflict potential, 90 interviews were performed in 3 communities in the study area, to assess wheter long term jaguar presence or movement through established corridors would be acceptable for the people living there. Key findings were that there was a negative impact of oil palm cultivation, especially for the species of focus. 48% of them were not recorded in palm oil habitats, and among them there were important jaguar prey species such as capybaras, spotted pacas, and collared peccaries, as well as pumas. Nevertheless, 65% of all species were recorded at least once in oil palm cultivated areas and the majority of them had improved RAIs at the edge of oil palm cultivated areas with secondary forest fragments or riparian vegetation, compared to oil palm habitats on their own. Jaguars were not frequently (or never for the female and her cubs) recorded in oil palm cultivated parts and their prey community was scarce, suggesting they can cross certain plantations but to survive and breed, they need more intact surrounding areas. Overall, it appears that small scale palm cultivation can be compatible with the persistence of some wildlife, if situated in a landscape that still includes less modified areas and if wildlife- favourable practices are adopted. Interview results indicate that attitudes are mostly positive, but 38% of respondents still considered jaguars problematic animals and there were some misconceptions especially amongst women. Hence, the area seems suitable, with the help of some awareness-raising activities, for establishing jaguar corridors, which will allow to guarantee this species long term connectivity.

Ai miei nonni, che vorrei con me indefinitamente y a los jaguares del Magdalena Medio.

A jaguar corridor can be “a cattle ranch, a citrus plantation, someone’s backyard – a place where jaguars can pass through safely and unharmed“ (Panthera).

Acknowledgements

This project and thesis were accomplished through the help of many people. First and foremost I would like to thank my parents, Enrica and Piero, my sister Chiara, and my grandparents Natalino and Noemi for their unconditional love and support, without which, none of what I do would be possible. Special thanks go to Cam for many reasons, for reviewing this manuscript, for getting his head around jaguarundi habitat use and Relative Abundance Indices, having studied history, and for getting rid of almost all my “italianisms”. Thanks also for all his support during the year and for dealing with my rollercoasters. Thanks for the best wines and weekends, and lastly, thanks for laughing with me about coo coos. A special thank goes to Panthera Colombia, for providing me with all kind of support, from anti-ameba medicines, to all transportation, accommodation and the possibility of using their camera traps, GPS and any other field equipment. I would like to deeply thank Dr. Esteban Payan, for a number of things. For his warm welcome and for everything I learnt from him, from jaguars and snakes to scientific paragraphs. Thanks also for his time and support as well as for good conversations, for dealing with all my questions, for instinctually trusting me from the very first conversation, and, finally, for giving me the amazing opportunity of joining Panthera and the Jaguar Corridor Initiative. Thanks to Dr. Marcus Rowcliffe for being such a good supervisor, for being reliable, for his time, input and patience at different phases of the project, his insights, his help with the analyses, and lastly, for revising this thesis. Thanks to all course supervisors, EJ Milner-Gulland, Marcus Rowcliffe, Colin Clubbe, and John Fa, for being there and for challenging us to think critically and transversally. I learnt a lot from all of you. Special thanks go to Carolina Soto for all her support, professional and personal, from picking me up at El Dorado on the first day to dinners and many other things. I would like to thank her also for her extreme professionalism and generosity, for teaching me elements of GIS and for producing all the maps I worked with as well as the ones reported in this manuscript. Special thanks also go to Carlos Marios Wagner for introducing me to the Middle Magdalena region and all of its wonderful people. Thanks for all his support in the field as well as his help during data analyses. Thanks also for giving me the opportunity of learning about camera trapping and the history of Colombia, together with unique moments and good conversations, and, finally for identifying the photos of birds.

I am extremely grateful to Cabildo Verde and James Murillo for all the local support and knowledge I received, and for finding me a home. Thanks to him also for his time, for the hours spent in “La Reserva” on weekends and for the nice time with his lovely wife Doriel and daughters Vanessa and Valentina. Thanks to Leidi, Mirella, Henry and Mario for offering me more than just a place to stay in the Middle Magdalena. Thanks for your warmth and for a feeling of home. Thanks to “El Loco” and Jaime for doing more than just transporting us, for helping with each camera trap and for being great and patient field guides. Thanks to Mildred, Johanna and Erika who guided me during the interviews, and who heard my script 40 times without losing their enthusiasm, and never stopped encouraging me. I am grateful also to “El Senor Mora” and “Don Julio” for welcoming me in their communities, for sharing with me their stories, knowledge, concerns, values, and hopes. Thanks to Caro, Loic and Lilli for being my family in Bogotá, for sharing the everyday, for giving me anchors when I was feeling lost, for welcoming me, and for those amazing dinners. Thanks to Diana for giving me her friendship straight away, for opening up to me and for being there. Thanks also for sharing with me your love for Colombia, for all those good conversations, for the maracuya margaritas, the Jardin Botanico, and much more. Many thanks to all the extended Panthera family for welcoming me, and because I learnt professionally and/or personally from all of you. And a special thanks to Angelica Benitez for always sharing her knowledge and for her help with Estimate S. My deepest gratitude go to all the women of my life, Alessandra, Anna, Annina, Barbara, Ceci, Chiara, Clem, Daniela, Elena, Enrica, Giulia, Giulia, Jenn, Lena, Luisa, Merel, Marina, Nina, Reka, Roberta, Silvia, and Whittney for being the friend and family of a lifetime. Colours would not be the same without you, and nothing ever would. Immense thanks go to Barbara and Clem, for walking next to me at all times, for being there at every step without being asked, and for teaching me a lot of what I know about friendship and its infinite value. Thanks to Madrid for its sun and its energy that changed my life forever, and thanks to Millie for making me fall in love with South American felids viscerally and irreversibly. Lastly, I would like to deeply thank Andy, Jack and Jenn, for making a difference every single day. Thanks for being the strongest support and the greatest companions through this Silwood adventure. Thanks for all your help, and above all, for our never-ending dinners, debates, and laughs that kept us sane and happy throughout the year. One additional thanks goes to Andy for his magical help with excel.

This project was entirely funded by Panthera Colombia.

1. Introduction

1.1 Problem statement

The jaguar, Panthera onca, is the only living representative of the genus Panthera found in the new world and is the largest cat existing in America (Nowell and Jackson 1996). Jaguars are flagship, umbrella and keystone species (Noss 1990, Caro et al. 2004, Soisalo and Cavalcanti 2006, Estes et al. 2011) with a strong current and past cultural value (Saunders 1998).

Jaguars are still abundant, but they are decreasing (Caso et al. 2008a), and they are estimated to occupy about 46% of their historical range (Sanderson et al. 2002). The species is considered Near Threatened due to habitat loss, caused by human population growth, agricultural expansion and economic growth; persecution; and poaching of its prey (Caso et al. 2008a). If these threats are not be mitigated jaguars overall will be qualified as Vulnerable in the near future (Caso et al. 2008a), as they already are in Colombia (Rodríguez-Mahecha et al. 2006).

It has been widely recognized that future of carnivores cannot rely on protected areas alone (Woodroffe 2001, Cardillo et al. 2003, Crooks et al. 2011), and the jaguar is no exception. Its conservation requires instead international, range-wide planning, and a landscape approach (Sanderson et al. 2002). This understanding has lead to range-wide priority setting and planning exercises. In the case of jaguars, experts have mapped 90 known existing population of jaguars with suitable habitat and prey base, Jaguar Conservation Units (JCUs) (Sanderson et al. 2002, Zeller 2007), and modeled probable corridors linking them (Rabinowtz and Zeller 2010). The resulting maps and knowledge were the foundation of the Panthera Jaguar Corridor Initiative, which aims at connecting jaguar populations from Mexico to Argentina (Fig.1.1).

Fig.1.1 Map of Panthera’s Jaguar Corridor Initiative. Corridors of concern are corridors less than 10km in width at any point along their length. The JCUs mapped were defined in 2006. Source: Panthera (2012).

This project unfolds within the Initiative. The success of the Jaguar Corridor Initiative, and, ultimately, of jaguar conservation strongly relies on understanding how jaguars use the remaining natural or degraded habitat within human-dominated landscapes, on the identification of the anthropogenic factors and land uses that are predominant in determining their habitat use, and finally on the adoption of landscape-level measures to effectively manage human-modified landscapes (Foster et al. 2010, Zeller et al. 2011).

Colombia is considered an area where efforts would significantly contribute to a range-wide jaguar conservation strategy, both because it is a critical link for jaguar connectivity between Central and South America, and because it contains some of the highest priority Jaguars Conservation Units and several corridors of concern, corridors less than 10km in width at any point along their length (Rabinowitz and Zeller 2010).

1.1.1 The Middle Magdalena region

Within Colombia, part of the region of Middle Magdalena (Fig. 1.2) became of great importance to Panthera Colombia and jaguar connectivity.

Its importance is due to its strategic position between two JCUs, San Lucas and Catatumbo (Fig. 1.1), and also because connectivity of the population of San Lucas and, ultimately, of Central American populations with the Amazon and Orinoquia populations is thought to

happen through this region (E. Payan, pers. comm.). This is because, although a direct corridor linking San Luca with the Orinoquia and Amazon populations was identified by the model and mapped, jaguars do not seem to use it due to the altitudinal barrier represented by the Andean Oriental mountains (E. Payan, pers. comm.) (Fig.1.1). The corridor was produced by the model because of its overall “least coast” and in theory jaguars can be found as high up as 2700m (Nowell and Jackson 1996), however they prefer low altitudes (Nowell and Jackson 1996, Zeller et al. 2011).

This region, even if heavily transformed by human activities, has also proven presence of jaguars (Panthera, unpublished data), well outside the areas and corridors identified by Rabinowitz and Zeller (2010). This has suggested a need to redefine a habitat connectivity strategy in the region and to re-establish corridors and areas actually used by jaguars.

Attitudes towards jaguars in this crucial region have not been assessed yet, and for a wide- ranging species with high-conflict potential to survive in an anthropogenic landscape, it is important to consider the human dimension (Marker et al. 2003). Understanding people’s perceptions towards jaguars will also finally inform the development of tailored initiatives and solutions.

Lastly, jaguars in the Middle Magdalena region, as in other areas of Colombia, have to face an additional threat: expanding oil palm cultivation. Colombia is currently the world's fifth largest palm oil exporter (Fedepalma 2012a). Furthermore, the industry is due to expand significantly by 2020 through increased production by a factor of 7 and the increase of cultivated land by 440% since 2000, according to Fedepalma, the Colombian plantation owners' association (Fedepalma 2012b). This poses a significant threat because palm oil plantations have been repeatedly proven to be extremely poor habitats for many vertebrate species (Maddox et al. 2007, Fitzherbert et al. 2008, Danielsen et al. 2009), including tigers (Panthera tigris) (Maddox et al. 2007), and to date no studies have investigated the impact of these plantation on jaguars or on other fauna of Colombia.

1.2 Project significance

Given the decline in number of jaguars, the increase of oil palm plantations, and the importance of the Middle Magdalena region for jaguar connectivity, this study will begin assessing, for the first time, what impact existing plantations have on the distribution and relative abundance of jaguars and other species detectable with camera traps. It will also produce the first data on the habitat use of these species across an oil palm plantation.

Findings will improve understanding of the extent to which these plantations represent a barrier for the movement of jaguars and to elaborate guidelines for oil-palm growers to reduce the impact of their plantations.

The project will also gather information on people’s attitudes towards this large cat, which will help identify if there is a potential for long term jaguars conservation in the area, if there are any misconceptions, and whether there is a need for awareness-raising activities. If so, results should inform the planning of these initiatives. Ultimately, this will make conservation efforts more effective.

Overall, understanding what passages jaguars actually use in this area and to what degree they are suitable, and safe, will help to manage the landscape accordingly, and to implement a better connectivity strategy in the Middle Magdalena region, an area which will particularly contribute to a range-wide jaguar conservation strategy.

1.3 Project objectives

Overall aim: contributing to the Jaguar Corridor Initiative through findings that will improve the planning of a habitat connectivity strategy for jaguars in the region of Middle Magdalena, Colombia.

Specific objectives with related research questions:

1. Begin evaluating the impact of existing oil palm plantations on species detectable with camera traps. Particular attention will be placed on jaguars, other Felidae species, their prey species and any species listed in IUCN or Regional Red Lists (all species of focus).

What species are found in oil palms cultivated areas in the region? How do oil palm cultivated areas compare with edge areas or oil palms free areas in term of species and/or their Relative Abundance Indices (RAIs)?

2. Investigate habitat use of jaguars and of their most common prey species (Central American agouti) in a context of an oil palm plantation and surroundings.

Are there any environmental features that are important in the habitat matrix for jaguar and agouti presence?

Findings deriving from both obj.1 and 2 will allow to begin compiling guidelines for oil palm growers and will improve the understanding of the extent to which oil palm plantations represent a threat for jaguar connectivity.

3. Describe the attitudes of three communities living in the study area towards jaguars to understand whether an established movement of jaguars in the area through corridors would be acceptable to the people living there.

What do people think of jaguars? Is long-term jaguar conservation in the area compatible with the views of local people? Are there misconceptions? Are awareness-raising activities needed?

Hypothesis:

The hypothesis regarding jaguars and oil palm cultivation is two-part. The first is that previous reports of jaguars using disturbed areas and degraded forest suggests that jaguars will be reported in the areas surrounding oil palm planted areas. The second part is that if jaguars are similar to tigers they will rarely or never be seen in oil palm cultivated parts. When considering other terrestrial mammals, similar results are expected. The negative impact of oil palm monoculture is therefore expected to be noticeable.

2. Background

The background consists of relevant literature, divided into the following sections: general knowledge on jaguars, reasons to conserve them, habitat use by jaguar, habitat use by other felids and common prey species, attitudes towards jaguars, and palm oil.

2.1 The jaguar

The jaguar (Fig 2.1), is the third-largest feline overall, after the tiger and the lion, it is solitary, and it is considered a top predator as well as an opportunistic hunter since its diet encompasses over 80 species (Nowell and Jackson 1996).

Fig. 2.1 Jaguar mother and two cubs photo-captured in the study area.

Its extent of occurrence ranges from Mexico to Argentina (8.75 million sq. Km) (Caso et al. 2008a), and while it has been estimated that 70% of jaguar range has a high probability of survival (Sanderson et al. 2002), ecological models found that most of it is actually of low suitability for jaguars (Torres et al. 2007). Jaguars density can vary from 7.5-8.8/100 km² in the Belizean Selva Maya rainforest (Silver et al. 2004) to 2.5/100 km² in unprotected areas in the Colombian Amazon basin (Payan 2009).

Jaguars are mostly nocturnal or crepuscular (Maffei et al. 2004, Harmsen et al. 2011). They were found to be diurnal in Brazilian Pantanal (Quigley and Crawshaw 1992), and active 24hrs in Peruvian rain forest (Emmons, 1987). These variations could be associated to the time of the day where their main prey (varying according to the area) are active (Quigley and Crawshaw 1992, Harmsen et al. 2011).

Large mammalian carnivores like jaguars are particularly vulnerable to habitat loss and fragmentation due to their biological traits, such as their body sizes, large area requirements, low densities and slow population growth rates (Cardillo et al. 2003). Furthermore, they are also vulnerable to persecution (Woodroffe 2001). Nowadays persecution of jaguars is predominantly in form of retaliatory kills following predation of livestock (Scogamillo et al. 2003, Polisar et al. 2003, Rabinowitz 2005), since the trade in jaguar skins, though considerable in the past (Ruiz-García et al. 2003), declined drastically after the mid-1970s, when CITES regulations were enforced (Nowell and Jackson 1996).

2.2 Why conserve jaguars?

The jaguar is considered a flagship and umbrella species (Caro et al. 2004, Soisalo and Cavalcanti 2006, Verissimo et al. 2012) as well as a keystone species (Paine 1969; Soisalo and Cavalcanti 2006, Estes et al. 2011). The latter is because apex predators, such as jaguars, through limiting populations of herbivorous mammals, play an important role in maintaining forest structure, which is ultimately connected to important ecosystem services such as water regulation (Estes et al. 2011). Though these effects are difficult to measure due to inherent ecosystems complexity and can be seen unequivocally only after apex predators have been lost (Estes et al. 2011), these authors were able to summarize evidence that confirm the role of top predators such as jaguars in shaping the ecosystem, using, among others, the example of Venezuelan forests on the islands of Lago Guri that showed reduced plant recruitment and survival in comparison to the mainland due to the disappearing of jaguars, cougars and harpy eagles, leading to a different herbivore assemblage.

In addition, jaguars have a strong cultural, spiritual and inspirational value demonstrated from their use as icons from sport cars to regional flags, banknotes and art works (Fig. 2.2) dating as far back as pre-Columbian civilizations in Central and South America (Saunders 1998).

Fig. 2.2 Cultural value of jaguars. From left to right: the flag of the department of the Amazonas in Colombia, a painting by Jorge Rajadell, and a “Moche Jaguar” (300AD), Larco Museum, Lima. Sources: from left to right, http://en.wikipedia.org/wiki/File:Flag_of_Amazonas_Department.svg, http://www.rajadellartgallery.com.ar/english/downloads/download_section.php?id_cat=21, and http://biology-forums.com/index.php?action=gallery;sa=view;id=3422

2.3 Measuring habitat use

The use of camera traps in investigating habitat use of medium to large terrestrial mammals has increased significantly in the last 5-10 years, especially when dealing with forest habitats and/or elusive species (Rowcliffe & Carbone 2008, Rovero et al. 2010). Most of the studies reported in the next two sections also employ camera traps data. Some environmental characteristics are recorded in situ (e.g. habitat type, canopy cover) at the camera locations, while others are extracted and quantified from satellite images. Studies reported consider the trap rates of species (or related measures) and compare them between habitat types (e.g. Di Bitetti et al. 2006) or use general linear models to investigate them in relation to environmental characteristics (e.g. Conde et al. 2010 and Davis et al. 2011). Correlations coefficents have also been used together with frequencies of locations with or without the species of interest (Foster et al. 2010, Rovero et al. 2010). However, these types of studies are limited by not addressing the issue of detectability (a species might be present in a site but not detected) (MacKenzie et al. 2002). Hence, other studies used occupancy models with environmental covariates to investigate habitat use with camera trap data (e.g. Negroes et al. 2010) or interview data (Zeller et al. 2011). Occupancy models were developed by MacKenzie et al. (2002), precisely to account for the issue of imperfect detection.

2.3.1 Jaguar habitat use

Jaguars occupy a variety of habitats including tropical and subtropical forest, pine-oak forest, unprotected forest buffer, disturbed forest, dense shrublands, agriclutural landscapes, wet grasslands, savannahs, shrubs, and pastures (Nowell and Jackson 1996, Monroy-Vilchis et al. 2009, Foster et al. 2010). However, they tend to favor forest over exposed areas with no vegetative cover such as pasture or agriculture (Schaller and Crawshaw 1980, Quigley and Crawshaw 1992, Michalski et al. 2006, Monroy-Vilchis et al. 2009, Zeller et al. 2011) and their presence is positively associated with water (Nowell and Jackson 1996, Michalski et al. 2006, Monroy-Vilchis et al. 2009) and canopy height (Conde et al. 2010, Davis et al. 2011). Male jaguars seem to be more tolerant to open areas than females (Conde et al. 2010, Foster et al. 2010).

Concerning the effect of roads, some contrasting results were found. Conde et al. (2010) reported that while roads had a negligible effect on males, females avoided them. Monroy- Vilchis et al. (2009) didn’ t distinguish between sexes and found that jaguars preferred distances between 3509 and 4377 m from roads. On the contrary, Davis et al. (2011) reported that jaguars presence was associated with roads within 200m but the roads in the study-site did

not hold heavy traffic, hence may actually provide a means of long distance travel and access to edge habitat where many prey are vulnerable.

Another factor affecting habitat use by jaguars is altitude. Jaguars usually inhabit environments at low altitudes, and avoid higher elevation areas and have not been found above 2,700 m in the Andes (Nowell and Jackson 1996, Zeller et al. 2011). Monroy-Vilchis et al. (2009) found that jaguars preferred altitudes higher than 1800m but this was attributed to the fact that human activity had displaced these large cats towards the higher parts of the study area, far from settlements.

Concerning the potential effects of hunting, fires or logging pressures on habitat preferences, Michalski and Peres (2005, 2007) found that no one of the three factors were significant for jaguars.

Regarding oil palm plantations, no studies have yet been carried out on jaguars. Tigers, while present in selectively logged and degraded forests (Linkie et al. 2008, Rayan and Wan Mohamad 2009) and even in unplanted areas of oil palm plantations, were never recorded in oil palm planted areas (Maddox et al. 2007).

For details on the studies reported (methods used, variables tested, significant conclusions) please refer to Appendix I.I.

2.3.2 Habitat use by other species

Habitat use will be described for the Felidae and prey species known to be common in the study area (Panthera, unplished data).

Pumas (Puma concolor) are found in a variety of habitats, from forests to lowland and montane deserts, pastures and croplands, though they prefer habitats with dense understorey vegetation and good forest quality and canopy cover (Michalski and Peres 2005, Davis et al. 2010). The puma can be considered more habitat generalist than the jaguar (Scognamillo et al. 2003, Foster et al. 2010). However, several studies have demonstrated that pumas tend to avoid areas of human activity, such as logging, hunting or travel (Ruth et al. 2003, Michalski and Peres 2005, Sweanor et al. 2008, Davis et al 2010 ). Pumas can be active throughout the day or can become predominantly diurnal or nocturnal (Harmsen et al. 2011, Paviolo et al. 2009). This could be due to temporal partitioning with jaguars, avoidance of human disturbance, or their prey habits (Emmons 1987, Paviolo et al. 2009, Harmsen et al. 2011).

Ocelots (Leopardus pardalis) inhabitat several habitat types, from all types of tropical forests, to

pine plantation, although with lower abundances (Lyra-Jorge et al. 2008, Di Bitetti et al. 2006). Ocelots also have high tolerance to seismic activities (Kolowski and Alonso 2010). They have been shown to be associated with well-watered habitats such as riparian forests and structurally closed habitats (Lopez-Gonzalez 2003, Di Bitetti et al. 2006). Ocelots are nocturnal animals (Crawshaw and Quigley 1989, Maffei et al. 2005, Di Bitetti et al. 2006).

Jaguarundis (Puma yagouarundi) occur in both closed and open habitats (provided some vegetative cover), from primary forests to grasslands and even pine plantations (Michalski et al. 2006, Nowell and Jackson 1996). They are considered more tolerant than other felids to human disturbance and degraded habitats (Michalski et al. 2006, Caso et al. 2008b). Given its diurnal nature, the jaguarundi is the Neotropical felid most easily seen (Caso et al. 2008b).

Central American agoutis (Dasyprocta punctuata) are found in forests, gardens and plantations (Emmons and Freer 1997), and can exploit from primary forests to highly disturbed habitats (Naughton-Treves et al. 2003). They are considered important seed dispersers for a number of tree species (Wright et al. 2007). Agoutis are diurnal with possible peaks of nocturnal activity (Lambert et al. 2009).

Among other prey species, spotted pacas (Cuniculus paca), although found in highly disturbed areas in some studies (e.g. Naughton-Treves et al. 2003), are recorded more frequently in densely forested habitats (Goulard et al. 2009, Trolle and Kéry 2005).

Lastly, the crab-eating raccoon (Procyon cancrivorous) is often associated with presence of water and better forest quality (Eisenberg 1990, Michalski and Peres 2005), suggesting that is less adaptable to human disturbance than the common raccoon (Reid and Helgen 2008).

2.4 Attitudes towards jaguars

Carnivores can easily generate negative attitudes among rural communities who are sharing their lands with them, and predators such as jaguars are also blamed for monetary losses following predation of livestock (Polisar et al. 2003, Palmeira et al. 2008).

Hence, it becomes crucial for carnivore conservation, including jaguars, to understand human perceptions to finally address potential misconceptions and conflicts (Marker et al. 2003). This has been done through surveys of local communities (semi-structured or structured interviews, and questionnaires) (Conforti and De Azevedo 2003, Rodrigues dos Santos et al. 2008, Figel et al. 2011).

Most studies report that the majority of people considers jaguars a positive presence and that

they should not be eliminated (Conforti and De Azevedo 2003, Rodrigues dos Santos et al. 2008, Figel et al. 2011). No differences between genders’ opinions were reported by Conforti and de Azevedo (2003), and having experienced a predation problem does not always affect respondents’ opinions (Zimmerman et al. 2005, Ávila-Nájera et al. 2011).

Despite of positive attitudes, people still think jaguars are problematic animals, mainly because they prey on livestock (Figel et al. 2011) but also because they can respresent a threat for human safety (Conforti and De Azevedo 2003, Rodrigues dos Santos et al. 2008). Perceptions of jaguars as a danger generally occur as a consequence of folklore and traditions (Wilson 2004) since only a minority of people had actually heard of an attack to humans (Rodrigues dos Santos et al. 2008).

Another issue is that most people seem not aware of the ecological role of jaguars in the ecosystem (Conforti and De Azevedo 2003) or of the effect of deforestation and habitat loss on wild populations (Ávila-Nájera et al. 2011).

While it is important to understand attitudes towards jaguars and large predators, it is also important to bear in mind that the relationship between attitudes and behaviors is not always powerful or straightforward as human behavior (e.g. jaguar killing) is influenced not only by personal attitudes but also by social norms and subjective norms (Marchini and Mc Donalds 2012). Therefore, investigating personal and collective norms that would make jaguar killing unacceptable from both a personal and a social point of view is also crucial (Marchini and Mc Donalds 2012).

2.5 Palm oil

Habitat loss driven by expansion and intensification of agriculture is currently considered the greatest threat to biodiversity (Tilman et al. 2001). Oil palm Elaeis guineensis already covers over 13.5 million hectares of tropical, high rainfall and lowland areas, which are natuarally occupied by tropical forests, the most biodiverse ecosystem on the planet (Fizhebert et al. 2008, Turner et al. 2008). In addition, palm oil production is increasing by 9% every year due to its highest yeld per ha among all vegetable oils (Butler and Laurance 2009) and to the growing demand for biofuels markets and for food production, especially in Indonesia, India, and China (Fizhebert et al. 2008).

Palm oil and its derivatives are indeed used under a variety of names in a wide spectrum of products (see Appendix I.II and I.III) from edible oils to ice creams, cosmetics, animal feed and biofuels (Bozzi 2007). It is estimated for example that one out of ten products in a UK

supermarket contains palm oil (Hickman 2006).

Currently 80% of palm oil is produced in Malaysia and Indonesia, but 410–570 million ha of currently forested land across Southeast Asia, Latin America and Central Africa are suitable for oil palm cultivation and might be converted as the demand rises (Fizhebert et al. 2008).

Colombia is listed as one of the world’s “megadiverse” countries, covering 0.7% of the planet but hosting ca. 10% of the planet’s biodiversity (Rodríguez-Mahecha. et al. 2006), and nowadays it is also the world's fifth producer of palm oil and the leading producer in Latin America (Bozzi 2007). A total of 300.000 hectares are cultivated among the departments of: Meta (1), Cesar (2), Santander (3), Magdalena (4), Nariño (5), Casanare (6), Bolívar (7), Cundinamarca (8), Chocó (9), and Norte de Santander (10) (Fedepalma 2012a) (Fig. 2.3).

Fig. 2.3 From left to right: departments of Colombia with oil palm cultivation (green), an oil palm plantation, and oil palm fruit. Sources (from left to right): Bozzi (2007), http://www.pinfosys.com.my/, and http://www.agricorner.com/palm-oil-climbs-to-28-month-high-on-prospects-for-higher-demand/

Palm oil accounts for 90% of the fats produced in Colombia. 60% of the overall palm oil produced is consumed in the country, while the remaining 40% is exported (Bozzi 2007), to the United Kingdom in the first place (Mingorance 2006). Oil palm growing is regarded as: “one of the country's most promising and important agricultural sectors and is at the heart of Colombia 's economic and social development” (Fedepalma 2012a). Furthermore, with a total of 3.5 millions hectares of suitable land for palm oil cultivation (Mingorance 2006), Fedepalma, the Colombian plantation owners' association, stated a 2020 vision of a seven-fold increase in production of palm oil and a 440% increase of cultivated land (Fedepalma 2012b).

This poses a concern for conservation because of existing evidence that oil palm plantation make poor habitats for the majority of species, compared not only to forests and degraded forest (Edwards et al. 2010, Peh et al. 2006, Fizhebert et al. 2008, Maddox et al. 2007, Danielsen et al. 2009) but even to other types of plantations such as rubber, coffee, and cocoa

(Edwards et al. 2010, Peh et al. 2006, Fizhebert et al. 2008). Indeed, only 15% across all taxa examined by Fizhebert et al. (2008) or 38% of the vertebrate groups considered by Danielsen et al. (2009) that were found in primary forests were also recorded in oil palm plantations.

Maddox et al. (2007) combined the use of camera traps, line transect surveys and radio collaring to investigate the impact of palm oil on tigers and large terrestrial mammals in an oil palm plantation concession in central Sumatra. Results were that 45% of the 38 species recorded in the landscape as a whole were detected at least once in the oil palm crop at some point and only 10% were regularly recorded in the crop, none of which has a high conservation value. Again, tigers, were never recorded in oil palm planted areas. Conversely, leopard cats (Prionailurus bengalensis) were regularly recorded in oil palm areas by Maddox et al. (2007) and Rajaratnam et al. (2007) found that this species even preferred oil palm habitats to forest fragments probably due to higher visibility and ease of movements resulting in greater hunting success. However, the cats still relied on forest fragments for resting and breeding.

In addition to direct impact on terrestrial above-ground biodiversity through habitat loss, palm oil monoculture also typically requires insecticides, rodenticides, herbicides and fertilizers that pollute soil and streams, ultimately affecting acquatic and soil biodiversity (Fizhebert et al. 2008). Hence, how to make palm oil a more wildlife friendly crop is an urgent issue and one to this point under-researched, given that only 1% of scientific publications regarding palm oil deal with its effect on biodiversity (Fizhebert et al. 2008).

Alternatively to wildlife-friendly agriculture, another option that has been proposed to reduce the impact of increasing crop production on biodiversity, is land sparing, which minimize the demand for farmland by maximising yelds. It is based on the concept that farmlands, regardless of practices adopted, host fewer species than relatively intact habitats, and this is especially true for species with high conservation value (Green et al. 2005). While there is evidence that land-sparing could allow more species to persist, maximising yelds usually also requires an increased used of pesticides or chemicals which would have a spill over effect on the sorrounding non-farmed areas. Furthermore, it is not guaranteed that non-cultivated areas would be left intact and not later converted to other non-favourable uses for biodiversity (Green et al. 2005).

Other concerns of oil palm monoculture deal with essential ecosystem services, such as carbon storage, that are significantly diminished in oil palm plantations (Butler and Laurance 2009).

Overall, the ecological impact of palm oil depends strictly on the amount of deforestation carried out to establish the plantation (Wilcove and Koh 2010) and the issue is that even when

legislation is in place to prevent forest loss, the revenue obtained through forest logging and timber production offsets the costs of establishing the plantation, making deforestation a strongly favourable option (Butler and Laurance 2009).

The impact of palm oil is not even limited to the ecological world (Mingorance 2006) since the establishment of oil palm plantations has already been associated with several crimes and human rights violations in Colombia as well as in the Middle Magdalena region itself, such as illegal and violent appropriation of land, massacres, and forced displacement (Mingorance 2006).

These concerns of both ecological and social nature have prompted the establishment of the Roundtable of Sustainable Palm Oil (RSPO) in 2004, providing certifications to palm oil producers who apply a series of principles and criteria involving both environmental and social sustainability. However, the overall efficacy of such certification schemes is still debated (Laurance et al. 2010) and in Colombia only one company is RSPO certified (RSPO 2012).

Regardless all of wildlife-friendly practices, land sparing, green markets, and certification schemes, to significantly reduce the rate of deforastation for oil palm cultivation the increasing demand for vegetable oils needs to be curbed as well (Koh and Lee 2012).

Despite its high impact, oil-palm agriculture has improved the livelihoods of many rural communities, thanks to improving their access to education, health, and infrustructures (Bozzi 2007). In conclusion, palm oil is a complex reality, and as Wilcow and Kow (2010) state, “there is no single best approach for dealing with the oil-palm crisis (…); a mixture of regulations, incentives, and disincentives targeted at all sectors of the oil-palm industry is necessary to protect (…) rapidly disappearing forests”.

3. Methods

3.1 Conceptual framework

The conceptual framework of the project, containing also the methods used to address the objectives and related research questions, is respresented in the following diagram (Fig. 3.1).

Fig. 3.1 Diagram representing the conceptual framework of the project (CTs=Camera Traps and PAs=Protected Areas)

3.2 Study area

The study was conducted within the Middle Magdalena region, which covers the central area of the Magdalena River basin, and in the municipality of Puerto Wilches, in a lowland area of ca. 60 sq. kilometers, situated to the west of the Magdalena River and to the north-east of the the laguna Cienaga de Paredes (Fig. 3.2). The study area and surroundings comprise secondary forests, shrubs, swamps, pastures, crops, oil palm plantations, and urban areas (Fig. 3.2 and 3.3a-b).

Fig. 3.2 Map of the study area, by Carolina Soto.

Although heavily transformed by human activities such as extensive livestock, oil extraction and expanding oil palm plantations, the study area has proven presence of jaguars as well as pumas, ocelots, jaguarundis, and prey species such as Central American agoutis, capybaras, pacas, opossums, nine-banded armadillos, and South American tapirs (Panthera, unpublished results).

Camera traps data included in this thesis were collected from a private property named: “El Hato”, 1800ha, where hunting is forbidden and oil palm growing is performed on a small scale. Six hundred and forty ha of the property are oil palm cultivated and managed by 25-30 workers. Management of the plantation includes enforcement of the law stating that at least

30m of riparian vegetation from any body of water should be left, as well as maintaining understory vegetation and vegetation growing on palm trunks. Working hours, and therefore maximum disturbance are from 6-7am to 3pm. A complete characterisation of the property, including chemicals and fertilizers used, was obtained through interviews to the plantation manager and manager assistant (see Appendix II.I).

Interviews were conducted in three rural communities sourrounding El Hato: “Cerrito”, “Campo Duro” and “Caño Limon” (Fig. 3.2 and 3.3d), which hold 82, 75 and 6 inhabited households respectively. Main activities in the communities involve fishing, agriculture, some livestock farming and manual labor in oil palm plantations. Level of infrustructure and technology varies among the communities, with the greatest level available at Campo Duro, slightly less so at Cerrito and considerably less so at Caño Limon.

Fig. 3.3 Vegetation types and some project activities. (a) Edge of secondary forest in El Hato, (b) oil palm cultivated area in El Hato, (c) retrieving camera trap data in El Hato, (d) conducting interviews in the community Cerrito. Photos a-b by V. Boron, c-d by C.M.Wagner.

3.3 Camera trapping

Camera-trap data have been collected since the end of January across “El Hato” an oil palm plantation concession, from 22 locations, covering an area of ca. 30 sq. km (Fig 3.2). The camera traps (CTs) (Cuddebacks, Bushnell Trophy and Panthera), triggered by movement, were both in oil palm cultivated areas and unplanted areas (secondary forests, pastures and swamps) (Fig.3.3a-b). These cameras were collected in mid-May, totaling 1190 camera trap nights, and were installed, one week later, in 25 locations in an area adjacent to the previous one and without oil palm cultivation (“Caño Limon”) (Fig. 3.2). Due to time constraints, only the dataset from the first session of camera trapping will be included in this thesis.

Jaguars are known to use trails (Silver et al. 2004), therefore cameras were placed on trails (2- 9m wide) to maximize jaguars encounters and/or at locations where jaguar sights or secondary signs had been reported from the plantation workers or local people, as by Silver et al. 2004, Maffei et al. 2004, Monroy-Vilchis et al. 2009. The distance of 1km between cameras was chosen to ensure that no individual home ranges of target species (Felidae sp.) would fall entirely between stations (Maffei et al. 2004). Cameras were placed at a height of 15-37 cm to ensure that smaller species would also be captured (Kelly 2008).

3.4 Habitat characteristics recording

Land cover classes in the study area and surroundings have been extracted and mapped from satellite images (ArcGIS). However the information available is from 2006 and considered too old for a fast-changing area such as the region of Middle Magdalena and not sufficiently detailed.

Therefore, the type of habitat was recorded in situ at camera stations and at 50m in all four directions (north, south, west and east). Habitat was classified as follows: secondary forest and edge of secondary forest bordering with pastures or roads “Forest + edge” (5 CTs), oil palm cultivated area (“Palm”) (6 CTs), edge between an oil palm cultivated area and a forest fragment (“Palm/Forest”) (7 CTs), and edge between an oil palm cultivated area and riparian vegetation (“Palm/Riparian”) (5 CTs). The last two habitat categories (“Palm/Forest” and “Palm/Riparian”) were also considered together as “Palm edge” (12 CTs). Canopy cover was assessed in situ, through subjective estimation and averaging of people estimates.

Distances from roads and settlements were not extracted from satellite images (for the same reasons stated above), and instead, RAIs of vehicles, people and livestock were used in the analyses as proxies for disturbance (as in Foster et al. 2010).

3.5 Interviews

90 semi-structured interviews to assess attitudes towards jaguars were conducted in “Caño Limon”, “Cerrito” and “Tierra Prometida”. Semi-structured interviews have been used successfully to gather information on attitudes of people towards large predators in previous studies (e.g. Conforti and Azevedo 2003).

Interviews were conducted with a random-sampling design aimed at covering half of the households in “Cerrito” (41 interviews) and “Campo Duro” (38 interviews), while interviewing all the people in “Tierra Prometida” (11 interviews), which can be considered a representative sample for the three communities (Newing et al. 2011). Only one person per household was interviewed in “Cerrito” and “Campo Duro”, alternating men and women, while in “Tierra Prometida”, due to limited number of inhabited households, all available people were interviewed.

Key aspects involved asking whether people considered jaguars problematic animals, whether they like to share their land with them, whether jaguars should be conserved and why, whether jaguars are more, less or equally common as before, and killing of jaguars. General background information such as age, occupation, number of kids, n. of years living in the area was also be recorded. People less than 15 years old or that had lived in the area less than 5 years were not interviewed, because they could be lacking perspective on present jaguar numbers compared to past ones or on jaguar killing in the past. For a full version of the interview script, refer to Appendix II.II.

3.6 Analyses

3.6.1 Camera trap and environmental data

RAIs, habitat comparisons and daily activity patterns: Relative Abundance Indices (RAIs) were calculated for each species and each habitat type, using the total number of independent capture events of that species in that habitat type divided by the number of trap-nights and expressed as records per 100 trap nights (Carbone et al. 2001, Kelly and Holub 2008). Independent capture events were defined defined as “ (1) consecutive photographs of recognizably different individuals of the same or different species, (2) consecutive photographs of individuals of the same species taken more than 30 min apart” (O’Brien et al. 2003).

These rules were not applied to calculate RAIs of people, vehicles, and livestock. This was because they were used as a proxy for disturbance, so amount of activity was considered of primary importance, regardless of the number of individuals involved.

While the debate continues on whether trapping rates can be used as an index of species density (Carbone et al. 2001, Tobler et al. 2008), they can certainly be used to investigate species occurrence in relation to habitat variables or to one another (Kelly and Holub 2008, Tobler et al. 2008). Therefore, RAIs of Felidae species, prey species and species listed in global or national red lists were calculated at each camera station and compared between habitat types using Kruskal Wallis or Mann Whitney tests (Payan 2009 and Rovero et al. 2010), since normality was not met. Post-hoc pairwise comparisons were performed adjusting the significance level to the number of comparisons, using Holm correction. Values of p<0.05 were considered significant (**) and values of 0.05≤p≤0.1 moderately significant (*). Since oil palm cultivation is expanding rapidly, and this expansion is favoured by decision makers at the national level, a slightly overestimation of its impact seems preferable to the risk of its underestimation. Spearman correlations were tested between RAIs of species and between RAIs of species and RAIs of vehicles, people and livestock. The statistical software used for the comparisons and correlations was SPSS, ver. 10.5.

Each photo included the exact time it was taken, which allowed the derivation of daily activity patterns of felids and the common prey species (photographed more than 10 times), by considering the percentages of photos taken per time interval of 2 hours. Six time intervals were in daylight (6am-6pm) and six during dark hours (6pm-6am).

Occupancy analyses: Occupancy analyses were performed for jaguars and the Central American agouti (most frequently recorded prey species at the study site). To account for imperfect detection and consequent understimation of occupancy, MacKenzie et al. (2002 and 2006) developed a model that estimates occupancy and detection probability based on repeated presence-absence surveys (detection histories) at multiple sites. In these models occupancy and detection probability can also be allowed to vary with covariates, entered using a logistic model, which allow to investigate species habitat use by assessing what environmental features (covariates) produce the best models. Here, the aim behind the use of occupancy models, more than producing the “best” estimates of occupancy and detection probability, is precisely to investigate habitat use.

Detection histories were constructed for each site (camera location) grouping 14 camera trap nights into one survey occasion (total of 8 survey occasions), and were imported in PRESENCE software (ver. 3.1) together with covariates. Occupancy models were run using a single-species single-season option and were designed assuming the following general hypotheses and site-specific covariates: 1) jaguar and agouti occupancy can be influenced by

the level of human disturbance (RAIs of people, vehicles and livestock); 2) jaguar and agouti occupancy can be influenced by habitat characteristics (type and/or canopy cover); 3) jaguar only: jaguar presence is determined by prey availability (RAIs of prey). Presence of water could not be tested alone as all of the 5 camera stations near the water were also at the edge of oil palm areas and are the ones classified as “Palm/Riparian”. It was therefore modeled as a categorical variable, i.e. “Palm/Riparian” habitat present vs. absent.

First, occupancy was estimated with the most simple model, where both occupancy and detection probability remained constant, ψ[.] p [.], then it was allowed to vary with each covariate considered singly (habitat type, canopy cover, RAI of people, RAI of vehicles, RAI of livestock, “Palm/Riparian” habitat present or absent, and RAI of prey for the jaguar only). Following this, occupancy was modeled with combinations of covariates that had produced models with deltaAIC<7 in the previous step. All models run were reported.

For sampling-occasion “season” was chosen as a covariate and modeled as a categorical variable, wet vs. dry (O’Connel et al. 2006).

Akaike Information Criterion (AIC) values were used to rank candidate models and Akaike’s weights (w) were used to interpret the relative importance of each model (Burnham and Anderson, 2002). Unless one model had a weight of 0.90, other models were considered when drawing inferences about the data (Burnham and Anderson 2002). Also, when there was a number of top ranked candidate models, i.e. deltaAIC<2 and similar AIC weights that were greater than 0.010, a model averaging technique was applied to estimate occupancy and detection probability from these multiple models (Burnham and Anderson, 2002).

3.6.2 Interviews data

Data from the interviews were mostly descriptive, Chi-squared tests were performed in SPSS (ver. 10.5) to assess whether gender or belonging to different communities influenced interviewees responses. Values of p<0.05 were considered significant.

4. Results

The results are divided in 3 main sections. The first one presents the results of the camera trapping (obj. 1-2) for all species and specifically for the species of focus. Then, it follows a section on the occupancy and habitat use analyses by the jaguar and the Central American agouti (obj. 2). Lastly, the third section contains the results of the interviews and addresses the attitudes towards jaguars (obj. 3).

4.1 Camera trapping

The total sampling effort across the 22 locations, from the end of January 2012 until mid-May of 2012 was of 1190 camera trap nights of which, 320 were in the habitat type “Forest + edge”, 329 in “Palm/Forest”, 175 in “Palm/Riparian”, and 370 in “Palm”. Independent capture events of wild species totaled 1686.

4.1.1 All species

A total of 47 species were detected, belonging to 20 orders (for a photo-inventory of the species detected refer to Appendix III). Mammals accounted for 21 (45%) of the 47 species detected and 7 orders, and birds for the remaining 26 species (55%) and 13 orders. Among mammals, 7 species were listed in IUCN or Regional Red Lists, while among birds only 2. No “Endangered” or “Critically Endangered” species were detected across the study area and Red Listed species were either “Vulnerable” or “Near Threatened” (Table 4.1).

Table 4.1 List of species recorded by CTs in study area and their IUCN and Regional Red List categories.

Scientific name Common name IUCN Red List Regional Red categorya list categoryb Birds Anseriformes Chauna chavaria Northern Screamer NT VU Cairina moschata Muscovy duck LC / Caprimulgiformes LC / Nyctidromus albicollis Pauraque LC / Charadriiformes Vanellus chilensis Southern lapwing LC / Ciconiiformes Agamia agami Agami heron VU / Bubulcus ibis Cattle egret LC / Cochlearius cochlearius Boat-billed heron LC / Egretta caerulea Little blue heron LC / Pilherodius pileatus Capped heron LC / Tigrisoma lineatum Rufescent tiger heron LC / Columbiformes

Patagioenas cayennensis Pale-vented pigeon LC / Coraciiformes Momotus momota Blue-crowned motmot LC / Cuculiformes Crotophaga ani Smooth-billed ani LC / Crotophaga major Greater ani LC / Falconiformes Geranospiza caerulescens Crane hawk LC / Buteo magnirostris Roadside Hawk LC / Cathartes aura Turkey vulture LC / Caracara cheriway Northern crested caracara LC / Milvago chimachima Yellow-headed caracara LC / Galliformes Ortalis columbiana Colombian cachalaca LC / Gruiformes Aramus guarauna Limpkin LC / Aramides cajaneus Grey-necked wood rail LC / Passeriformes Furnarius leucopus Pale-legged horner LC / Oryzoborus angolensis Chestnut-bellied seed-finch LC / Piciformes Galbula ruficauda Rufous-tailed jacamar LC / Tinamiformes Crypturellus soui Little tinamou LC /

Mammals Carnivora Cerdocyon thous Crab-eating fox LC / Leopardus pardalis Ocelot LC NT Panthera onca Jaguar NT VU Puma concolor Puma LC NT Puma yagouaroundi Jaguarundi LC / Eira barbara Tayra LC / Lontra longicaudis Neotropical otter DD VU Procyon cancrivorus Crab-eating raccoon LC / Cetartiodactyla Pecari tajacu Collared peccary LC / Chiroptera Unknown Bat Didelphimorphia Didelphis marsupialis Common opossum LC / Metachirus nudicaudatus Brown four-eyed opossum LC / Pilosa Myrmecophaga tridactyla Giant anteater VU VU Tamandua tetradactyla Lesser anteater LC / Primates Aotus griseimembra Grey-handed night monkey VU VU Cebus albifrons White-fronted capuchin LC NT Rodentia Hydrochoerus hydrochaeris Capybara LC / Cuniculus paca Spotted paca LC / Dasyprocta punctata Central American agouti LC / Sciurus granatensis Red-tailed squirrel LC / Unknown Mouse a IUCN 2012. b Renjifo et al. 2002 and Rodriguez-Mahecha et al. 2006.

The species accumulation curve (Fig 4.1a) doesn’t level off, suggesting more species could have been detected with a larger sampling effort. However, when considering mammals and birds separately it appears that the increase in the total number of species is caused by an increase in birds species detected, whereas the mammal accumulation curve does level off after ca. 800 trap nights (Fig 4.1b).

Fig 4.1 Species accumulation curves. (a) All species, (b) mammals and birds considered separately.

The most common species recorded were the crab-eating fox (Cerdocyon thous), and the Central American agouti (Dasyprocta punctata) for mammals, and the paraque (Nyctidromus albicollis) and the grey-necked wood rail (Aramides cajaneus) for birds (Table 4.2). Overall, the most common and most ubiquitous species was the crab-eating fox, present at 20 out of the 22 camera stations (91%).

Table 4.2 Number of independent capture events and RAIs per 100 trap nights, in each habitat type and overall, for all species detected. Highlighted rows contain “Species of Focus” (Section 4.1.2). Bold species name indicate species are listed in IUCN or Regional Red Lists.

Forest + edge Palm edge Palm Total Capture Capture Capture Capture RAI RAI RAI RAI events events events events Mammals Aotus griseimembra 0 0 0 0 18 4.83 18 1.51 Bat 0 0 1 0.20 0 0 1 0.08 Cebus albifrons 1 0.31 8 1.59 0 0 9 0.76 Cerdocyon thous 244 76.25 150 29.76 187 50.13 581 48.82 Cuniculus paca 7 2.19 5 0.99 0 0 12 1.01 Dasyprocta punctata 137 42.81 65 12.90 4 1.07 206 17.31 Didelphis marsupialis 0 0 0 0 1 0.27 1 0.08 Eira barbara 2 0.63 1 0.20 1 0.27 4 0.34 Hydrochoerus hydrochaeris 1 0.31 0 0 0 0 1 0.08 Leopardus pardalis 11 3.44 6 1.19 7 1.88 24 2.02

Lontra longicaudis 0 0 0 0 1 0.27 1 0.08 Metachirus nudicaudatus 1 0.31 0 0 2 0.54 3 0.25 Mouse 0 0 1 0.20 1 0.27 2 0.17 Myrmecophaga tridactyla 0 0 1 0.20 1 0.27 2 0.17 Panthera onca 9 2.81 38 7.54 4 1.07 51 4.29 Pecari tajacu 0 0 1 0.20 0 0 1 0.08 Procyon cancrivorus 5 1.56 8 1.59 31 8.31 44 3.70 Puma concolor 12 3.75 2 0.40 0 0 14 1.18 Puma yagouaroundi 5 1.56 4 0.79 5 1.34 14 1.18 Sciurus granatensis 2 0.63 13 2.58 15 4.02 30 2.52 Tamandua tetradactyla 0 0 2 0.40 0 0 2 0.17

Birds Agamia agami 0 0 1 0.20 0 0 1 0.08 Aramides cajaneus 76 23.75 76 15.08 16 4.29 168 14.12 Aramus guarauna 1 0.31 8 1.59 1 0.27 10 0.84 Bubulcus ibis 5 1.56 1 0.20 3 0.80 9 0.76 Buteo magnirostris 1 0.31 1 0.20 0 0 2 0.17 Cairina moschata 0 0 0 0 1 0.27 1 0.08 Caracara cheriway 0 0 1 0.20 5 1.34 6 0.50 Cathartes aura 0 0 3 0.60 12 3.22 15 1.26 Chauna chavaria 0 0 3 0.60 0 0 3 0.25 Cochlearius cochlearius 0 0 1 0.20 0 0 1 0.08 Crotophaga ani 5 1.56 9 1.79 2 0.54 16 1.34 Crotophaga major 1 0.31 1 0.20 1 0.27 3 0.25 Crypturellus soui 2 0.63 0 0 0 0 2 0.17 Egretta caerulea 1 0.31 0 0 0 0 1 0.08 Furnarius leucopus 0 0 0 0 6 1.61 6 0.50 Galbula ruficauda 1 0.31 0 0 0 0 1 0.08 Geranospiza caerulescens 1 0.31 0 0 0 0 1 0.08 Milvago chimachima 1 0.31 6 1.19 29 7.77 36 3.03 Momotus momota 3 0.94 0 0 0 0 3 0.25 Nyctidromus albicollis 6 1.88 141 27.98 65 17.43 212 17.82 Ortalis columbiana 5 1.56 28 5.56 46 12.33 79 6.64 Oryzoborus angolensis 1 0.31 0 0 0 0 1 0.08 Patagioenas cayennensis 13 4.06 43 8.53 23 6.17 79 6.64 Pilherodius pileatus 1 0.31 0 0 1 0.27 2 0.08 Tigrisoma lineatum 3 0.94 2 0.40 0 0 5 0.42 Vanellus chilensis 0 0 0 0 1 0.27 1 0.08

Average Mammals 34 6.50 15 2.89 20 3.55 49 4.09 Average Birds 7 1.47 12 2.39 13 2.12 25 2.07 Average all species 18 3.60 13 2.60 16 2.74 35 2.95 T0T. Photos 564 631 491 1686

The three habitat types had identical number of species (31), although a different species composition. Out of the 18 species never recorded in “Palm” 7 were mammals (33%) and 11

birds (67%). Of the species never recorded in “Palm”, the capybara (Hydrochoerus hydrochaeris), the blue-crowned motmot (Momotus momota), the little blue heron (Egretta caerulea), and the roufous-tailed jamacar (Galbula ruficauda) were never recorded in “Palm/edge” either. However, almost all these species were only photographed once (Table 4.2).

Species with the highest RAI in “Forest + edge” compared to the other habitat types include the puma (Puma concolor), the spotted paca (Cuniculus paca), the rufuscent tiger heron (Tigrisoma lineatum), the smooth-billed ani (Crotophaga ani), and the cattle egret (Bubulculus ibis) (Table 4.2).

Of the 31 species found in palm, only 8 (26%) had higher RAIs in this habitat than in others, 3 mammals, the crab-eating raccoon (Procyon cancrivorous), the grey-handed night monkey (Aotus griseiembra), and the red tailed squirrel (Sciurus granatensis), and 5 birds, the southern lapwing (Vanellus chilensis), the Colombian cachalaca (Ortalis colombiana), the yellow-headed caracara (Milvago chimachima), the pale-legged horner (Furnarius leucopus), and the turkey vulture (Cathartes aura) (Table 4.2).

Overall, RAIs averaged across all species are similar in different habitat types, while the average RAI of Mammals in “Forest + edge” is higher than the RAI of Mammals in “Palm” or “Palm Edge” (Table 4.2). However, these differences were not statistically tested, as in this project, particular focus was given only to Felidae species, their prey species and any species listed in IUCN or Regional Red Lists.

4.1.2 Species of focus

Felidae species, their prey species and any species listed in IUCN or Regional Red Lists totaled 17 species (highlighted rows, Table 4.2). The species most frequently recorded was the Central American agouti, followed by the jaguar, and the crab-eating raccoon (Fig. 4.2).

Fig. 4.2 Species of focus most frequently photo-captured. From left to right: Central American agouti (Dasyprocta punctuata), jaguar (Panthera onca), and crab-eating raccoon (Procyon cancrivorous).

The Central American agouti although common, was found at 10 out of the 22 stations (45%), while crab-eating raccoons were more ubiquitous and recorded at 15 stations (68%). Jaguar was detected at 12 camera stations (54%). When considering the other felids (Fig. 4.3a), pumas were found in 5 stations (23%) and ocelots and jaguarundis at 10 (45%) and 9 (41%) stations respectively. There was a overlap between the stations where cats were recorded, especially between jaguars and pumas, and ocelots and jaguarundis (Fig. 4.3b).

(a)

(b) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 16 17 18 19 20 21 22 23 Jaguar Puma Ocelot Jaguarundi

8 (48%) of the 17 “Species of Focus” were never recorded in oil palm habitat and of the 9 species recorded in “Palm” only two (crab-eating raccoon and grey-handed night monkey) seem to prefer it (higher RAIs) to the other habitats (Table 4.2). Jaguars were recorded in palm areas with much lower RAI than in the “palm edge” habitat and the female jaguar with the cubs was never detected in the “Palm” habitat.

At first, to understand species preferences in regards to palm-edge habitat, the two types of palm-edge habitats were considered separately as “Palm/Forest” and “Palm/Riparian”. Some species were recorded in only one of the two habitat types, with the majority of them in “Palm/Forest” (Fig. 4.4). RAIs were compared for the species recorded in both types, however the differences were never significant (Mann Whitney test, p>0.1), therefore the two habitats were again combined and treated as “Palm edge”.

Agamia agami Chauna chavaria

Aotus griseimembra Cebus albifrons Cuniculus paca Dasyprocta punctata Didelphis marsupialis Hydrochoerus hydrochaeris Leopardus pardalis Metachirus nudicaudatus Species ofSpecies focus Myrmecophaga tridactyla Palm/Riparian Panthera onca Palm/Forest Pecari tajacu Procyon cancrivorus Puma concolor Puma yagouaroundi Tamandua tetradactyla

0.00 5.00 10.00 15.00 20.00 25.00 RAIs Fig. 4.4 RAIs for the Species of Focus, calculated directly from the number of independent capture events and the number of trap nights for each palm edge habitat, and expressed as records per 100 trap nights.

Some species were recorded in all three habitat types (“Forest + edge”, “Palm edge” and “Palm”), some species only in two and some only in one (Table 4.2 and Fig. 4.4). For the species present in all three habitat types (the jaguarundi, the crab-eating raccoon, the jaguar and the ocelot) RAIs were compared using Kruskal Wallis and no significant differences were found (p>0.1). Mann Whittney tests were performed for species recorded in only two of the three habitat types (the white-fronted capuchin, the puma and the spotted paca), and a significant difference was found for the puma (U=7.0, p=0.012**).

Cumulative RAIs of all “Species of Focus” and of “Felids and prey species” were also considered (Fig. 4.5) and a significant differences were found between habitat types (H=6.06, d.f.=2, p=0.048** and H=5.14, d.f=2, p=0.067* respectively).

All Species of Focus TR Palm TR palm edge

Felidae + prey TR Forest + edge

0.00 10.00 20.00 30.00 40.00 50.00 60.00 70.00

RAIs

Fig. 4.5 Cumulative RAIs for the Species of Focus and Felidae and prey species RAIs for the Species of Focus, calculated directly from the number of independent capture events and the number of trap nights for each habitat type, and expressed as records per 100 trap nights.

Post-hoc pairwise comparisons (Mann Whitney tests) where carried out between “Palm” vs. “Forest + edge” and “Palm” vs. “Palm edge”. For “Felids and prey species” there was a significant difference between “Palm” and “Forest + edge” (U=3.0, p=0.050*) and for all “Species of focus” there were significant differences both between “Palm” and “Forest + edge” (U=3.0, p=0.050*) and between “Palm” and “Palm edge” (U=27.0, p=0.061*). Levels of significance were adjusted following Holm correction, significance for the lower value of p between the two post-hoc comparisons was set at 0.1/N=0.05 (where N=2 is the number of comparisons), while significance for the higher value of p was set at 0.1/(N-1)=0.1.

No correlations were found between RAIs of species or between RAIs of species and RAIs of vehicles, people and livestock, with the exception of the RAI of vehicles that was positively correlated with the RAI of jaguars (ρs=0.471, p=0.004**) and negatively correlated with the

RAI of Central American agoutis (ρs=-0.592, p=0.027**).

Daily activity patterns were considered for the four felids species and the three most common prey species, the Central American agouti, the crab-eating raccoon and the spotted paca. Crab- eating raccoons and spotted pacas were markedly nocturnal with all photo-captures taking place from 6pm to 6am and 4am respectively. Jaguars, pumas and ocelot were predominantly crepuscular and nocturnal (majority of photo-captures occurred from 4pm to 6am), and displayed moderate activity early in the morning (6am-8am). Central American agoutis and jaguarundis were diurnal and crepuscular, they were photographed the most from 6am to 8pm and from 8am to 8pm respectively (Fig. 4.6). It is worth noticing that for all nocturnal cats, jaguars, pumas and ocelots, the few photos (2, 1 and 3 respectively) taken at times that overlap with the plantation working hours (7am-3pm), were not taken by cameras located in the oil palm cultivated areas (“Palm”).

Fig. 4.6 Daily activity patterns of felids and most common prey species (N=number of independent capture events (y axes) per time interval, expressed as % of the total number.

4.2 Occupancy and habitat use analyses for the jaguar and the agouti

206 Central American agouti and 51 jaguar independent capture events were obtained in the study area from 10 and 12 camera stations, producing naive occupancy estimates of 0.60 for the jaguar and of 0.50 for the agouti. Two out of the 22 camera stations were not included in the occupancy analyses because the cameras were mostly non functional.

The simplest model with constant occupancy (ψ(.)) and constant detection probability (p(.))had poor support in for both the jaguar and the agouti (Models J10 and A6, Table 4.3).

Table 4.3 Occupancy models for the jaguar (J1-J17) and the central American agouti (A1-A8). Covariates “vehicles”, “people”, “livestock”, and ”prey” were included as RAIs. “PR” stand for “Palm/Riparian” habitat, hence for vicinity of water.

Model ID MODELS AIC deltaAIC AICweight Likelihood N.Par. J1 ψ (vehicles+can.cov.), p(.) 105.20 0 0.220 1 4 J2 ψ (vehicles), p(.) 105.86 0.66 0.155 0.719 3 J3 ψ (vehicles+can.cov.+people), p(.) 106.60 1.40 0.107 0.497 5 J4 ψ (vehicles+prey), p(.) 106.66 1.46 0.104 0.482 4 J5 ψ (vehicles+people), p(.) 106.76 1.56 0.099 0.458 4 J6 ψ (vehicles+can.cov.+prey), p(.) 107.19 1.99 0.080 0.370 5 J7 ψ (vehicles+livestock), p(.) 107.49 2.29 0.069 0.318 4 J8 ψ (vehicles+prey+people+ can.cov.), p(.) 108.59 3.39 0.034 0.184 6 J9 ψ (vehicles+can.cov+PR), p(.) 109.13 3.93 0.030 0.140 6

J10 ψ (.), p(.) 109.36 4.16 0.027 0.125 2 J11 ψ (vehicles+PR), p(.) 109.50 4.30 0.025 0.116 5 J12 ψ (can.cov.), p(.) 111.28 6.08 0.010 0.048 3 J13 ψ (PR), p(.) 111.32 6.12 0.010 0.047 4 J14 ψ (prey), p(.) 111.35 6.15 0.010 0.046 3 J15 ψ (people), p(.) 111.36 6.16 0.010 0.046 3 J16 ψ (livestock), p(.) 112.58 7.38 0.005 0.025 3 J17 ψ (habitat), p(.) 114.71 9.51 0.002 0.009 6

A1 ψ (can.cov.+vehicles), p(.) 85.94 0 0.868 1 4 A2 ψ (can.cov), p(.) 90.07 4.13 0.110 0.127 3 A3 ψ (vehicles), p(.) 93.29 7.35 0.022 0.025 3 A4 ψ (PR), p(.) 103.26 17.32 0.000 0.000 4 A5 ψ (livestock), p(.) 105.25 19.31 0.000 0.000 3 A6 ψ (.), p(.) 105.37 19.43 0.000 0.000 2 A7 ψ (people),p(.) 107.10 21.16 0.000 0.000 3 A8 ψ (habitat),p(.) 107.20 21.26 0.000 0.000 6

When detection probability was allowed to vary upon the season (wet vs. dry), it resulted in worse models for the jaguars (increase of the AIC of 1.69-2.00 units from models with the same parameters and p constant) and comparable models for the agouti (increase or decrease of the AIC of 0.03-0.43 units), therefore models with p(season) were excluded from the analyses and p was always considered constant (p(.)).

The best ranked models for the jaguar (deltaAIC<2 and AICweight >0.10) (Burnham and Anderson, 2002 and Linkie et al. 2007) were the ones where occupancy was allowed to vary with RAI of vehicles and canopy cover, or vehicles alone, or vehicles, canopy cover and people, or vehicles and prey (Model J1-J4, Table 4.3). All of these responses were positive.

For the Central American agouti there was one best model and it was the one where occupancy varied with canopy cover and RAI of vehicles (Model A1, Table 4.3). In this case, the RAI of vehicles had a negative effect on agouti occupancy while canopy cover had a positive effect.

For the jaguar, since, no single model emerged as the top ranked model, i.e. AICweight>0.90, weighted averages of ψ and p were produced from models with a weight of at least 0.10 and a deltaAIC<2 (Models J1-4, Table 4.3). The averaged estimate of occupancy was equal to 0.65, while the averaged estimate of the probability of detecting a jaguar if present was of 0.53.

For the agouti, even if the AICweight of the best model (Model A1, Table 3.3) was not greater than 0.90, it was definitely close (0.87) and significantly higher than the one of the second best model. Hence the model (A1, Table 4.3) was considered a single top ranked model and

occupancy and probability of detection were estimated from this model, at 0.60 and 0.63 respectively.

4.3 Attitudes towards jaguars

90 interviews were conducted across the three communities, 41 in Cerrito (21 women and 20 men), 38 in Campo Duro (20 women and 18 men) and 11 in Caño Limon (4 women and 7 men). Overall respondents were 50% women and 50% men.

The average age of respondents was 37.0±13.3 (SD) (16-66). The people interviewed had lived in the area from 5-64 years, for an average of 19.1±12.3 (SD) years and had on average 3 children (0-11). Most women were housewives (77.8%), some (13.3%) practiced agriculture and a minority (8.9%) worked in a shop. Men’s activities displayed more variation, with 37.8% of men working in oil palm plantations, 15.6% practicing agriculture, 8.9% fishing, 2.2% farming livestock, 8.9% living off a combination of two of the activities already mentioned, and the remaining 31.1% living of “everything, depending on availability and current needs”.

100% of people stated that there are still jaguars in the area, and 83% would like/likes to see them in the wild. There was a significant effect of gender on responses (χ²=9.356, d.f.=2, p=0.009**), indeed while no man said “No” and only 4% said “From far away”, 16% of women said “No” and 11% “From far away”. In Campo Duro 7 respondents (18%) had seen a jaguar, in Cerrito 15 (37%) and in Tierra Prometida 7 (64%). Sightings occurred across the 24hours, both in light and dark hours.

When people were asked whether they liked to share their lands with jaguars, responses were different between communities (χ²=7.002, d.f.=3, p=0.030*) (Fig. 4.7). Among the 11 interviewees who gave a negative response, 9 (82%) were from Cerrito and the remaining 2 (18%) from Campo Duro. In Caño Limon all respondents liked to share their lands with jaguars.

100%

80% Fig. 4.7 Communities’ preference 60% (“Yes” or “No”) regarding the 40% No sharing of lands with jaguars, expressed as percentages of “Yes” 20% Yes and “No” per each community. 0% Campo Duro Cerrito Caño Limon

Again, when people where asked whether they thought that jaguars are problematic animals, there was a significant effect of the community on the response (χ²=9.566, d.f.=4, p=0.048*). Cerrito was the village were the highest percentage of people (55%) considered that jaguars are or could be problematic animals (Fig. 4.8).

100% 80% Fig. 4.8 Communities’ opinion on whether jaguars are problematic 60% No animals (“Yes”, “No” and “Could 40% Could be be”), expressed per each community 20% Yes as percentages of respondents giving 0% each answer. Campo Duro Cerrito Caño Limon

Overall, across all communities, the reasons for considering jaguars problematic animals involved in order of decreasing importance, possible conflicts with livestock, attacks on people and livestock, attacks on people, and a perceived aggressiveness of jaguars that frightens people (Fig. 4.9).

Attacks on people Fig. 4.9 Reasons for 12% considering jaguars Attacks on people and livestock 9% problematic animals, 40% Perceived aggressiveness expressed as percentages Conflicts with livestock of respondents giving 29% each reason.

Most of the people (65%) considering the conflict with livestock as the main problem were men, while most of the ones (80%) considering jaguars’ aggressiveness as the main issue were women.

When interviewees were asked if they thought jaguars could attack people without being provoked, 24% responded “Yes”, 7% “Maybe” and 69% “No”. Gender influenced the response (χ²=9.580, d.f.=2, p=0.008**) and 70% of the people that answered “Yes” or

“Maybe” were women (Fig. 4.10). Only 11 out of the 21 respondents who believed that jaguars could attack people said that they had actually heard about such an attack and they were unable to define specifically the location and the circumstances.

100% Fig. 4.10 Genders’ opinion on 80% whether jaguar can attack people without being provoked (“Yes”, 60% No “No” and “Maybe”), expressed per 40% Maybe each gender as percentages of Yes respondents giving each answer. 20%

0% Women Men

However, despite jaguars possibly being problematic animals and possibly attacking people, 95% of all respondents think that is important to conserve them and 96% of people would like to have tourists coming to the areas to enjoy wildlife. People’s reasons for conserving jaguars are various: the fact that they are declining, their belonging to nature, their beauty, their non- aggressiveness, the possibility for future generations of knowing jaguars, the role of jaguars in their habiats, and religious beliefs (Fig. 4.11).

10% Decline 24% Belonging to nature Fig. 4.11 Reasons for conserving jaguars, expressed as percentages of 10% Beauty respondents giving each reason (this Future knowledge was an open question, however 12% Non-aggressiveness respondents only gave one reason) 18% Importance for habitat 17% Religious reasons

People were also asked whether they believed that jaguars were greater, less or the same in number than before. Overall, 39% believed that jaguars are greater, 2% that they are the same and the remaining 59% that they are less. Cerrito had the highest percentage of respondents answering that jaguars have increased in number (51%), Caño Limon had the lowest (20%) and

Campo Duro an intermediate value of 32%. The effect of gender on responses was almost significant (χ²=5.860, d.f.=2, p=0.053) and more women than men thought that jaguars have increased and vice versa (Fig 4.12).

100% Fig. 4.12 Genders’ opinion on 80% whether jaguar have increased 60% (“More”) or decreased (“Less”) in Less number, expressed per each gender 40% as percentages of respondents giving More each answer. 20%

0% Women Men

The reasons given for the decrease in jaguar numbers were in order of importance: habitat loss (29%), kills (24%), a combination of habitat loss and kills (20%), and jaguar displacement to other areas (10%). 17% did not give a specific reason and only commented that they saw them less than before. The awareness of habitat loss as a reason for jaguar decline was greatest in Campo Duro (50%) and lowest in Cerrito (21%), while the awareness of killing of jaguars as a cause for their decline was relatively high in Campo Duro and Cerrito, 58% and 68% respectively, and low in Caño Limon, 9%.

Reasons for jaguars increase in number included that they are reproducing (43%), that they are not killed anymore (52%), and a combination of the two factors (5%). Interestingly, the majority of the 15 respondents who believed jaguars are increasing in number because they are reproducing are women (73%) and, also, from Cerrito (60%).

Respondents were also asked about killing of jaguars. Overall, 12 people (13%), 6 from Campo Duro and 6 from Cerrito, answered that jaguars are still killed in the area, the method of killing is with guns and the main reason involves conflict with livestock. Only 2 people, 1 from Campo Duro and one from Cerrito, said jaguars were killed in the area within the last year.

In regards to kills of jaguars in the past the majority of people (73%) stated that few jaguars were killed in the past (<5), and 15% answered: “None”. Concerning kills of jaguars in the surrounding areas, slightly more people, 18 (20%) believed jaguars are still killed, of these, 3 were from Campo Duro (accounting for 9% of the people interviewed in the community), 9 were from Cerrito (33% of the total people interviewed in the community) and 6 were from Caño Limon (67% of the total people interviewed in the community).

5. Discussion

Jaguars are declining, and although species-based conservation has been criticized due to its constraints (Mace et al. 2007), conserving jaguars that are both umbrella and keystone species also allows to conserve lower trophic levels and the integrity of functional ecosystems as a whole (Estes et al. 2011). This study had developed with three specific objectives concerning (1) the impact of oil palm plantations on jaguars and other species detectable with CTs, (2) the habitat use of jaguars and Central American agoutis across an oil palm plantation concession, and (3) people attitudes towards jaguars. Hence, the discussion will begin with three sections that reflect both the result sections and the objectives. The following section includes insights from the results placed in the context of the original research questions and project objectives. Finally, the concluding section contains some key messages, implications for conservation, and recommendations.

5.1 Camera trapping and impact of oil palm cultivation

The property where camera trapping was conducted, although certainly human transformed, also contained some well conserved areas. This is because oil palm cultivation is practiced at a small scale since two thirds of the land cannot be cultivated for topographic reasons, leaving a varied habitat mosaic. Furthermore, hunting is forbidden in the property, riparian vegetation is left untouched for at least 30m from any body of water, understory vegetation in the oil palm cultivated areas is not cleared and chemicals for sanitary control are used only ad hoc. All these aspects make this concession a best case scenario in terms of oil palm cultivation. Hence, while results have the limitation of not being directly transferrable to larger and “worse” plantations, finding an impact of oil palm cultivation in this study, even at this scale, represents a clear and alarming result.

The total number of species recorded (47) was slightly higher than the one found by Payan (2009) in the Colombian Amazon or by Tobler et al. (2008) in the Amazonian moist forest of Peru, despite the sampling effort being lower in this study, indicating that partially degraded habitats are not necessarily impoverished in the number of species. However this should be proved by considering the total number of species present in the general area where the study took place. The data from the camera traps currently running in the area adjacent to the oil palm plantation concession, which is free of oil palm cultivation, will provide valuable information in this regard and will also improve the understanding of which between land

sparing and biodiversity-friendly agriculture could be the best option for mitigating the impact of oil palm cultivation on wildlife.

When considering mammals solely, the number of species (21) was lower than the one found by Tobler et al. (2008) or by Payan (2009), while comparable to the one recorded in the Colombian eastern Llanos (Díaz-Pulido et al. 2010). The leveling of the mammals’ accumulation curve indicates that even with a larger sampling effort no more mammal species were likely to be detected.

Mammal community was dominated by rodents and carnivores, and among them, the crab- eating fox and the Central American agouti were predominant. Both these species are known to be resilient and adaptable to certain degrees of habitat degradation (Maffei and Taber 2003, Naughton-Treves 2003). South American tapirs (Tapirus terrestris) and nine-banded armadillos (Dasypus novemcinctus), though known to be present in the general area (Panthera, unpublished results), were never recorded in the specific study area, which could be because both species, and especially tapirs, can be sensitive to human disturbance (Cullen et al. 2000, Hill et al. 2003). No species of deer were detected and even collared peccaries, known to be present along human disturbance gradients (Altrichter & Broaglio 2004), were detected at an incredibly low relative abundance. This situation could be symptomatic of an unbalanced mammal community, and not necessarily in terms of total number of species, but more in terms of composition and relative species abundance. However, a pleasant and unexpected surprise was recording giant anteaters and grey-hand night monkeys, given the rarity of both species (IUCN 2012).

Though carnivores, and, specifically, jaguars are present and relatively common, the prey community for jaguars seems impoverished, since, even if considered prey generalists (Nowell and Jackson 1996), they are known to rely on, armadillos, peccaries, and deer (Novack et al. 2005, Weckel et al. 2006, Foster et al. 2009), which are almost all absent here. This suggests that while jaguars use and cross this area they likely rely on more intact surrounding areas to survive. Another possibility could be that jaguars are preying on smaller species such as the crab-eating raccoon, the agouti and the paca, and supplementing their diet with domestic animals, as found by Foster et al. (2009). However, this is unlikely because even if jaguar activity pattern overlaps with crab-eating raccoons and pacas (both nocturnal species) there is an evident temporal partitioning between jaguars and agouti, the most frequently observed prey species. In addition, jaguar attacks on livestock in the oil palm plantation concession have not occurred in the last few years. Regardless, as this study did not perform any diet/scat

analyses, it cannot rigorously address to what extent the available prey community in the study area is sufficient for jaguars.

5.1.1 Species and habitat types

Overall, the ratio of species detected in oil palm cultivated areas to that overall (31/47) was high and definitely greater than that found by Maddox et al. (2007), but this can be attributed to the greater size of oil palm cultivated areas in Maddox et al. (2007). However, the species that had higher RAIs in oil palm cultivated areas were a minority, and among them only two were Red Listed species, the grey hand night monkey and the giant anteater, and the others were species with low conservation value such as the red-tailed squirrel and the crab-eating raccoon.

The Colombian cachalaca also preferred oil palm planted areas to the other habitat types, and photographic evidence show these birds eating palm fruits which could be an encouraging result since this species has an important role in maintaining the integrity of forest ecosystems as a seed disperser (Erard and Sabatier 1994). However, almost half of the “species of focus” were not even recorded once in palm oil habitats, and among them there were important jaguar prey species such as capybaras, spotted pacas and collared peccaries, as well as pumas. Jaguars were infrequently recorded in the palm habitat, and the mother with cubs never. Furthermore, even agoutis, photographed frequently across the study area, showed a preference for forested habitats and were barely recorded in “Palm”. Hence, the conservation value of oil palm cultivated area for jaguars and in general, although not null, it appears limited, and their negative impact noticeable, which supports the original hypothesis.

In palm edge habitats the situation improves, especially for the species of focus (cumulative RAI significantly higher). This shows that maintaining both riparian vegetation and, even more so, unplanted fragments of secondary forest (depending on the species) can be a valuable tool to make oil palm plantation more wildlife friendly as suggested by Fitzherbert et al. (2008) and Koh et al. 2009. These could act “stepping stones” for dispersal of species able to move across oil palm dominated areas (Wilcowe and Koh 2010). In addition, mantaining forested areas could also favour oil palm growers by containing the spread of deseases affecting palms.

Jaguars were found to tolerate oil palm edge habitat considerably well and to a greater extent than “Palm” habitat on its own. While their preference for the “Palm/Riparian” habitat is likely due to its vicinity to water, in accordance with previous studies (Nowell and Jackson 1996, Michalski et al. 2006, Monroy-Vilchis et al. 2009), they were also frequently recorded also

in the “Palm/Forest” habitat. Jaguars’ tolerance to palm edge habitat is in accordance with the original hypothesis, since these large cats have already been recorded in other types of disturbed habitats (Monroy-Vilchis et al. 2009, Foster et al. 2010). Furthermore, as jaguars were almost exclusively nocturnal in this study, the temporal overlap of human disturbance (plantation workers and vehicles) and jaguar activity was minimal. Overall jaguars seem to be more tolerant to oil palm cultivation than tigers that were never present in oil palm planted areas (Maddox et al. 2007).

While palm edge habitat had improved presence also for spotted pacas and pumas when compared to the “Palm” habitat (in which both species were never recorded), it showed significantly lower presence for pumas and lower presence for spotted pacas when compared to the “Forest + edge” habitat. Pacas were recorded more frequently in more densely forested habitats also by Goulard et al. (2009) and Trolle and Kéry (2005). Results regarding pumas suggest that they are less tolerant than jaguars to oil palm cultivation. While some literature considers pumas more habitat generalists than jaguars (Scognamillo et al. 2003, Silveira 2004), other authors, such as Foster et al. (2010) found them to be less tolerant than jaguars to human disturbance. Furthermore, they can be active during all 24 hours (Harmsen et al. 2011) and their predominant nocturnal activity at this site, could be their strategy to avoid disturbance, as found by Paviolo et al. (2009).

Another species that benefited from the palm/edge habitat in comparison to the “Palm” habitat was the Central American agouti, showing that although they are found in highly disturbed habitats (Naughton-Treves et al. 2003), their tolerance to oil palm areas might be limited.

Conversely, both ocelots and jaguarundis were found in “palm” more than in palm edge habitat and only slightly less (especially for the jaguarundi) than in forested or “forest + edge” areas, suggesting that they are tolerant species, in accordance with previous research (Di Bitetti et al. 2006, Caso et al. 2008b, Koloski and Alonso 2010). Their high RAIs in “Palm”, could also be due to a high visibility and prey availability in this habitat, made up of rodents, such as squirrels and mouse (the latter probably under-detected due to its size). These findings are in accordance to what found for for the leopard cat (Maddox et al. 2007 and Rajaratnam et al. 2007).

Overall, a factor that should be considered is that only 1 camera out of the 5 located in the “forest + edge” habitat, was actually in the forest. Hence, the “best” habitat type in the study is

already a disturbed habitat and therefore differences between the “Palm” habitat and actual secondary or primary forest areas should be expected to be greater and should be investigated.

5.2 Habitat use and occupancy analyses

The habitat use by jaguars and Central American agoutis, the most commonly recorded prey species, was investigated with occupancy models containing environmental covariates.

The fact that the best models for jaguars included the RAI of vehicles and that it had a positive association with jaguar occupancy might be because jaguars like to use the roads that are also more accessible to vehicles for their long distance movements. Jaguars are known to use trails (Silver et al. 2004), however their presence in relation to roads has shown contrasting tendencies (Monroy-Vilchis et al. 2009, Conde et al. 2010, Davis et al. 2011). In this case, given traffic was only diurnal and jaguars nocturnal, is reasonable to think that these roads were favoured for movements also because undisturbed at times were jaguar were active. Covariates such as canopy cover and the RAI of prey species as a whole also produced good models but only in combination with vehicles. Jaguars association with good canopy cover has been documented (Monroy-Vilchis et al. 2009, Zeller et al. 2011), however, models considering only canopy cover resulted in being poor models. Prey availability alone, was also a poor predictor of jaguar habitat selection.

In the best model for the Central American agouti RAI of vehicles had a negative effect on agouti occurrence and canopy cover a positive one. Since agoutis are diurnal animals and were found to be almost 100% diurnal in this study as well, it is reasonable to conclude that their presence can be affected by the traffic taking place from 7am to 3pm across the plantation, and that they rely on remaining forested areas with better canopy cover, in accordance with their much higher RAI in forested areas than in palm areas.

A possible limitation is that trail type or width were not considered in the analyses, however since all trails were wide (2-9m) their effect on detectability might be limited but should be investigated.

Both jaguar and agouti findings regarding vehicle disturbance highlight the importance for oil palm plantations to maintain working hours that do not extend over the 24hrs.

In conclusion, the results from the models seem to suggest that even if occupancy rates were relatively high for both species (ψjaguar =0.65 and ψagouti =0.60) there were differing spatial trends for jaguars and agoutis. These findings corroborate the other elements of evidence collected (temporal partitioning between jaguars and agoutis and poor prey availability in the

concession) in suggesting that jaguars cross this area as part of their movements, but not necessarily to hunt. Hence, while it can be said that small scale oil palm cultivation, if adopting favorable practices, might constitute a limited threat for jaguar connectivity, it is likely insufficient for their survival.

5.3 People attitudes towards jaguars

Overall, in this area most of the people believe it is important to conserve jaguars, and like to share their lands with them, which is an encoraging result in itself. However, one limitation of the interviews was that they were performed in communities where only a small percentage of people practiced livestock farming, rather than in cattle ranches, where more conflicts between people and jaguars are usually likely to occur, and attitudes could be worse.

Even if most people living in these three communities believe in the importance of conserving jaguars, and like to share their lands with them, 38% still thought jaguars are problematic animals, due to the conflict with livestock, their aggressiveness, and their perceived attacks to humans and livestock. Similar results were found by Conforti and De Azevedo (2003) and Ávila-Nájera et al. (2011). This is especially true in the community Cerrito which also holded the highest amount of interviewees who do not like to share their lands with jaguars as well as the highest number of people with the misconception that jaguars are increasing in number and the lowest percentage of respondents aware of habitat loss.

These results are probably due to a combination of factors. The community Caño Limon has grown to be extremely motivated in regard to nature conservation probably because of its isolation and direct dependency on its services. Instead, the community of Campo Duro, while much less dependent on nature than Caño Limon, has received environmental education from an oil extraction company, which could have increased people knowledge on local wildlife and threats. Cerrito is not in such direct contact with nature as Caño Limon and the awareness- raising activities in the community have been strongly focused around manatees and aquatic ecosystems. Both aspects, as well as community-specific cultural heritages, could explain a weaker knowledge and stronger reticence towards jaguars of the respondents living in Cerrito.

Significant differences did not emerge only between communities but also between respondents of different genders, in contrast to results by Conforti and De Azevedo (2003). Interestingly, more women than men tend to see jaguar aggressiveness as the main reason for considering jaguars problematic animals, and also more women believed jaguars could attack people without being provoked. Unlike lions or tigers, jaguars have never been reported to kill

a human (Conforti and Azevedo 2003). Also in this study only one of the people who claimed to have heard of such an attack was able to clearly specify the circumstances or any detail. Hence, it is likely that this perception of jaguars being aggressive is due to a percieved risk fueled by tradition, and orally transmitted stories. Finally, more women than men also thought that jaguars have increased in numbers and many of them gave as a reason that “they are reproducing”.

Again, the reasons behind these differences are various. Women are naturally less strong then men hence could be more fearful of jaguars. Nevertheless, it is also commonly believed that jaguars like to attack women, especially when they are pregnant, inducing a greater fear in women than in men. Lastly, because of the lifestyle women hold in these rural communities of Colombia (almost 80% of the interviewed ones are housewives), they tend to be more disconnected from reality than men are and this could increase their misconceptions.

Regardless of gender or community of belonging, only a minority (10%) of people, when asked why conserving jaguars, were aware of their role on the ecosystem, as found by Conforti and De Azevedo (2003). Another issue is that only one third of people seemed aware of habitat loss and respondents tend to have a restricted and localized view of the issue, believing jaguars are decreasing in the area but not overall, since they are just moving somewhere else where forests are not disappearing. The widespread lack of education could be responsible for these beliefs, and although the level of education was not assessed in this study, conversations with community leaders indicate that most people only have primary school.

Concerning jaguar killing, positive results were found. So, overall, results could suggest that the area is suitable for jaguar long term conservation and that agreements for the implementation of jaguar corridors could be achieved. However attitudes towards jaguar should be assessed across cattle ranches in the area and some awareness-raising initiatives would be beneficial in the community of Cerrito and especially targeting women.

5.4 Evidence from results in relation to project objectives and research questions

Project objectives, related research questions and project findings are summarized in Table 5.1.

Table 5.1 Summary of project’s findings in relation to initial objectives and research questions.

Objectives Research questions Findings 1. Begin evaluating the A. What species are found in oil A. 66% of species overall and 52% of the species impact of existing oil palm palms cultivated areas in the of focus were recorded in oil palm cultivated plantations on species region? areas at least once. Jaguars were recorded detectable with camera traps, infrequently and the mother with cubs never. and in particular on jaguars, Impoverished prey community for jaguars in the other Felidae species, their study area and even more so in oil palm planted prey species and any species areas which did not host pacas, peccaries, listed in IUCN or Regional capybaras as well as pumas. Red Lists (all species of focus). B. How do oil palm cultivated B. Overall, same number of species in “Palm”, areas compare with palm-edge “Palm edge” or “Forest + edge”, but different areas or oil palms free areas in species composition. Generally lower RAIs in term of species and/or their “Palm”, especially for the species of focus RAIs? (significant difference). Only two Red-listed species preferred “Palm” habitat to the other

habitat types.

Overall: noticeable negative impact of oil palm cultivation and limited potential for jaguar conservation (survival and breeding), and even lower potential for pumas conservation.

2. Investigate habitat use of A. Are there any environmental A. The presence of both species was positively jaguars and of their most features that are important in associated with canopy cover. Jaguars favoured common prey species in a the habitat matrix for jaguar locations also with higher RAI of vehicles, context of an oil palm and agouti presence? suggesting they use them for their long distance plantation and surroundings. movements. On the contrary, agoutis avoided locations with higher RAI of vehicles.

3. Describe the attitudes A. I. What do people think of A. I. 95% of people think is important to towards jaguars of three jaguars? conserve jaguars and 88% like to share their communities living in the lands with them. However 38% see them as study area to understand problematic animals due to possible conflicts whether an established with livestock and a perceived aggressiveness. movement of jaguars in the Cerrito was the community with the worst area through corridors would attitudes. be acceptable to the people II. Are there misconceptions? II. Few people are aware of jaguar ecological living there. role, or habitat loss. 40% of people believe jaguars are increasing in number. There was more misconception regarding jaguars among women than men.

B. Is long-term jaguar conservation B. People overall had positive attitudes, and 98% in the area compatible with the reported that jaguars are not killed in the area views of local people? anymore, suggesting the conservation of jaguars is compatible with people views.

C. Are awareness-raising activities C. Awareness activities would be beneficial to needed? address overall low environmental and jaguar knowledge. Cerrito should be prioritized as well as women.

5.5 Conclusion and suggestions

The expansion of the palm oil industry in Colombia is inevitable and also an opporunity for a further development of the country, hence efforts should be placed in reducing its ecological impact. Key messages of this research are that there was a negative impact of oil palm cultivation, especially in terms of RAIs of species and especially for the species of focus. Nevertheless, more than 60% of the species were recorded at least once in “Palm” habitat and the majority of species had improved RAIs in the “Palm edge” habitats compared to “Palm”. Jaguars were found in the study area, although were not frequently recorded (or never for the female and her cubs) in oil palm cultivated parts. In addition the prey community for jaguars was scarce, suggesting they can cross certain plantations but to survive and breed, they still need more intact surrounding areas with greater prey availability. Overall, it appears that small scale palm cultivation can be compatible with the persistence of some wildlife (resident or transient) if the following recommendations are applied: fragments of forest and riparian and understory vegetation are maintained, plantation working hours do not extend over the 24 hours, the use of chemicals is localized and the plantation is situated in a varied landscape that still includes less human modified areas and oil palms free areas. The latter is particular important, suggesting that the planning for the establishment or expansion of oil palm plantation should happen at the landscape level.

Other suggestions for making oil palm cultivation a more wildlife-friendly crop could involve the increase of incentives for RSPO or wildlife-friendly certifications through the establishment of green markets. Bateman et al. (2010) report for example that consumers in England are willing to pay a premium price for products containing “tiger-friendly” palm oil. The complementary approach (consumers pressuring both manifactureres and retailers to not use palm oil produced at the expenses of biodiversity) could also contribute to change producers’ behaviour (Bateman et al. 2010). Another option that has been proposed is for NGOs to buy small tracts of existing oil palm plantations. The revenue would first offset the cost of buying and then would allow to buy forested land to establish a network of private reserves for biodiversity conservation (Koh and Wilcowe 2007). Lasly, reducing consumers’ demand is extremely important as well, since the pressure to convert tropical forests into soy bean or oil palm cultivation could be halved by a decrease of vegetable fat consumption of 25g/day (=large serving of French fries) in the EU and US (Koh and Lee 2012).

In regard to land sparing, further results of this project will improve the understanding on whether more species are found in areas that are completely free of oil palm cultivation, and/or it they are more abundant. However, overall, since few areas of intact habitat are left in this region, land sparing might not be a viable option for the Middle Magdalena, but should still be considered for other regions and larger plantations, that might be too large and industrial to host jaguars and other species, regardless of the adoption of favourable practices.

Again, because this represents a best case scenario for oil palm cultivation, further research should be placed in investigating the effect of bigger and more industrial plantations managed under different circumstances and practices. This would allow to assess what features characterize the plantations were jaguars (and other species) are found. Such findings could be used to produce guidelines for oil palms growers to either establish and manage plantations that are more jaguar/wildlife-friendly or intensify their yields and adopt a land sparing strategy. Research should also assess if there are threshold dimensions of plantations, above which, jaguars crossing does not occur anymore. Finally, jaguar diet in a context of oil palm cultivation should also be studied.

From a human dimension, the area seems mostly suitable, with the help of some awareness- raising activities, for reaching agreements over the establishment of jaguar corridors, which will ultimately guarantee jaguars long term survival and connectivity.

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Appendix I

I.I Jaguar habitat use

Study Techniques Analyses Variables tested Significant results (Jaguar) Carnivore co-existence 47 camera stations in Gneralized linear models, % canopy cover, canopy height (m), canopy height, roads within and habitat use in the a grid at intervals of assuming a negative % understory cover, Log tree density, 200 m, small bird TSc large Mountain Pine Ridge c. 1.5–3.0km along binomial error Road width, River length, Road mammal TS Forest Reserve, Belize roads and trails distribution, with counts length, Distance to the Macal River, (Davis et al. 2011). 74 days of jaguar, puma, ocelot or Human TS, Small bird TS, Large bird Note: TS=Trap Success approaximately. grey fox captures at each TS, Small mammal TS, Medium station as the response. mammal TS, Large mammal TS. Habitat Use by Sympatric Camera traps at 178 Frequencies of locations Sex, habitat type, trail width, distance Males: present with the Jaguars and Pumas locations, totaling ca with and without big cats to water, distance to human highest frequency in pastures Across a Gradient of 22,000 trap-nights were compared between settlement, activity of jaguars of and up to 438m frm forest. Human Disturbance in over 3yr (2004– habitat (chi-square)A two- opposite sex, distances to the forest Trail width (+), female Belize (Foster et al. 2006). The distance factor general linear block, disturbed forest, savannah, activity (+), distance from 2010). between neighboring model (GLM) with pasture, plantation and major roads. pastures and plantations (+) stations averaged 0.8 repeated measures on Female: present more in km (SD = 0.8 km, N the random factor camera disturbed than in protected = 178). station forest, not over 178m from forest. Distance from water (- ) distance from forest (+), human development (+) Integrating Occupancy 71 6x6 km sampling Occupancy analyses using Habitat covariates included Occupancy: higher elevation modeling and interview units, 4-6 interviews PRESENCE (v3.0; proportion of forest, grassland, areas (-), areas with no data for corridor per unit to gather Hines, 2010) and habitat types (agriculture/ shrub, vegetative cover (-). identification: A case detection/non-detec- ArcView. open areas (bare ground and Detection probability: is study for jaguars in tion data on jaguar pastures), early-stage forest re- higher in lower elevation Nicaragua (Zeller et al. and several prey growth, wetland, and water), mean areas (-), inteerviewee survey 2011). species. elevation of a sampling unit and effort (+) and the proportion distance to protected areas. of water (+). Cougar and jaguar habitat Structured habitat use index (HUI) vegetation type (pine-oak forest, oak pine-oak forest habitats (+) , use and activity patterns interviews, for each variale evaluated. forest, tropical deciduous forest, altitudes higher than 1800 in central Mexico signs (tracks and expected frequency introduced grassland and agriculture), (+), distances between 3509 (Monroy-Vilchis et al. scats), and (Poisson distribution)- distance from settlements, distance and 4377 m from roads (+), 2009). 22 camera-traps. observed frequecy. from roads, distance from very steep between 2329 and 4650 m slopes (exceeding 50%) (six equal from settlements (+), and intervals), and altitude (six 300-m distances to very steep slopes altitude intervals). between 1048 and 2059 m (+) Sex matters: Modeling GPS collars were Generalized linear model Sex, land cover (tall forest, short Lowest AIC: models with male and female habitat programmed to (GLM) to infer the forest, swamp, secondary vegetation, interactions between sex and differences for jaguar record animal probability of jaguar and agriculture-cattle, which included road distance (S:R) and conservation (Conde et positions at varying occupancy as a function low-intensity, non-mechanized cattle between gender and land al. 2010). time intervals over of sex and environmental ranching and agricultural land), cover (S:L). Males and 2–12 months. variables. Distance to roads, distance to females: tall forests (+), 3 females population centers and human swamps and secondary 3 males population densities were not growth (-), agriculture-cattle (- included due to their strong ). Females: short and tall correlation with road proximity forests(+), proximity to roads (-). Males: short forest (-) Using occupancy models 119 camera traps Occupancy models Distance to the nearest water course, Distance from water (-) to investigate space over 3 months, distance and percentage of dense Dense habitat (+) partitioning between two distance of 3.5 km habitat (forest and scrublands as sympatric large between cameras. opposed to open grasslands and predators, the jaguar and swamps) in a 1750m buffer around puma in central Brazil the camera trap. (Sollman et al. 2011). Anthropogenic Interviews. T tests patch size Land cover classes (closed- canopy forest habitat quality (+), determinants of primate Patch occupancy data occupied vs unoccupied forest, semi-open forest, open- burn severity (+), patch size and carnivore local on the midsized to canopy forest, disturbed forest, (+), age of isolate patch (-). extinctions in a large-bodied highly disturbed forest, managed and fragmented forest vertebrate fauna of Step-down logistic unmanaged pasture, recent clear-cuts, landscape of southern each patch, including regression models bare ground, and open water), Amazonia (Michalski and five primate and 14 patch connectivity, spatial extent and Peres, 2005). carnivore species. intensity of hunting, logging and fires. Human–wildlife conflicts Visits and interviews Correlates of predation Land cover classes: (close and open Predation by jaguars affected in a fragmented to 236 cattle ranches. canopy forest, lightly and higly by bovine herd size (+) and Amazonian forest Validation of General linear models disturbed forest, managed and the proportion of forest area landscape: determinants occurrence of felids unmanaged pasture, bare ground and within a 5km buffer (+), of large felid depredation and predation events openwater), dist. to main town, age distance from the nearest on livestock (Michalski et through line-transect of the ranch, proportion of forest riparian corridor (-) al. 2006). censuses and a in concentric rings, cattle herd size, photo-trapping. staff ratio, dist. to riparian corridors.

I.II Different names for palm oil and its derivatives

Vegetable Oil, Vegetable Fat, Sodium Laureth Sulfate, Sodium Lauryl Sulfate, Sodium Dodecyl Sulphate (SDS or NaDS), Palm Kernel, Palm Oil Kernel, Palm Fruit Oil, Palmate, Palmitate, Palmolein, Glyceryl Stearate, Stearic Acid, Elaeis Guineensis, Palmitic Acid, Palmitoyl oxostearamide, Palmitoyl tetrapeptide-3, Steareth -2, Steareth -20, Sodium Kernelate, Sodium Palm Kernelate, Sodium Lauryl Lactylate/Sulphate, Sodium Lauryl Sulfoacetate, Hyrated Palm Glycerides, Sodium Isostearoyl Lactylaye, Cetyl Palmitate, Octyl Palmitate, Cetyl Alcohol, Palmityl Alchohol.

I.III The oil palm production chain: stages and products (Bozzi 2007) RBD: Refined Bleached Deodorised.

Appendix II

II.I Characterisation of the oil palm plantation concession “El Hato”

GENERAL INFORMATIONS Department: Santander Municipality: Puerto Wilches Vereda: Pampas Coordinates of the “Reception/main office” : 18N 0819845 0631912 Dimension: 1800ha Oil palm planted area: 640ha N. of palms: 90.000 palms. Ca. 141 palms/ha. Distance between palms: ca. 8.3 m N. of workers: 25-30, paid per amount of fruits collected. Working hours: 6am to 3 pm. People living in the property: ca. 12 Year of establishment of oil palm cultivation: 2000 Growth of palms: 40-50cm/yr Year of first harvest of fruit: 2004 End of production: after 25 years, palms are 25m tall. Too tall to harvest. Current plans of expansion: NO expansions planned, topographic reasons. Productivity: 140-180 tons of fruits in 2 weeks. ca. 4160 tons per year . ca. 6,5 tons/yr/ha Processing of fruit: locally. Uses: Edible items, no bio-fuels. Previous use of property: pastures for livestock farming. Conservation of riparian vegetation : 30 m along streams and ponds (lagunas? Cienagas?) Hunting: prohibited in the property

OIL PALM PLANTATION MANAGEMENT Fertilizers: • NITRAX-S 28-4-0-6 (s) Ammonium Nitrogen (NH4N) 17.5%, Nitric Nitrogen 10.5%; Assimilable Phosphorus (P2O5) 4%; Sulfur (S) 6% • NUTRIMON 13-6-23-6 Ammonium Nitrogen (NH4N) 8.3%, Nitric Nitrogen 4.7%; Assimilable Phosphorus (P2O5) 6%; Potassium soluble in water (K2O) 23%; Magnesium (MgO) 6%; Sulfur (S) 3%; Boron (B) 0.25%. Frequency: Twice a year when raining, for a better soil penetration.

Diseases and control: Note: Insecticides and fungicides are only delivered when/if needed and only on ill palms, mainly for PC control. There is no systematic/general use of these chemicals.

• PC-Bud rot complex Caused by: complex of fungi. Outbreak: 2009 % Palms affected: 10-12%, of which 40% are in good state. Limited outbreak of PC is possibly due to two reasons: forests surrounding the plantation act as a barrier to impede PC transmission from other plantations, and the soil is highly compact, which decreases water absorption from the palms, which results, in turn, in less humidity and therefore less favorable conditions for the fungi. Control: SEVIN PO(o 80?) WP, insecticide; RIDOMIL GOLD, fungicide; VITAVAK, prevent fungi from colonizing; FOSFITO, fungicide. • Rhynchophorous palmarum Insect, eats buds previously rotten by fungi (PC) and ultimately cause the death of the palm. Control: traps with insecticide and natural through ants whose nests are in the vegetation of the trunk and in the understory vegetation. • Cephaloleia vagelineata Insect, eats buds, natural control. • Salagasa ocrhacea It targets roots, chemical control, last present 6 yrs ago. • Strategus atoeus Beetle, eats leaves, natural control. • Ossiphanes cassina Butterfly larvae, eat leaves, natural control (ants).

Management of palms and lots • Palm trunks Cleared only at the base, vegetation left all along • Palm pruning Once a year • Lots clearing Understory vegetation cleared in plates at the base of the palms, manually with machine, and naturally by leaving palms branches on the ground at the base of the trunk. The use of GLIFOSATO stopped in 2009 due to its high levels of toxicity. Major paths are cleared alternatively. Same path is cleared every-other time of fruit collection in the lot (24 to 30 days) to allow vegetation to re-grow. • Collection of fruit Manual: Rotation scheme covering all lots, lasting 12-15 days, then back to the 1st lot. Animals: mules, buffaloes, and horses collect fruit from the lots and transport them to the main path where a truck collects them.

Vehicles/means of transport: • Main roads and collection trails All means of transport: truck, tractors, motorbikes, horses, mules, buffaloes. Main roads, transit occurs daily. Collection trails, less traffic, related only to that lot (collection/clearing/pruning) • In the lots Tractors, people walking, horses, buffaloes (fruit collection/clearing/pruning/treatment of ill palms)

Sources of information: Mr. Avelardo, help-manager of the plantation, semi-structured interview on 15/05/2012 & Mr. Alfredo, manager of the plantation, semi-structured interview on 12/06/2012

II.II Interviews script

Attitudes towards jaguars

Panthera Colombia – Imperial College London May-June 2012

1. Basic information

1.1 Interview # ____ 1.2 Date:______1.3 Departament: __Santander__ 1.4 Municipality:______1.5 District______1.6 GPS Coordinates: ______

1.7 Gender: Man (__) Women (__)

1.8 Wherewere you born?______1.9 How old are you? ______

1.10 How long have you been living in the area?______

1.11 How many children do you have? ______

1.12 How many people depend on you? ___ Adults ___ Kids (<15 yrs old)

1.13 What is your occupation? Livestock (__) Agriculture (__) Fishing (__) Palm (__) Home(__) Shop (__) Other: ______

2. Attitudes towards jaguars

2.1 Would you like to see a jaguar in the wild? Yes (__) From far away (__) Don’t care (__) No (__)

2.2 Are there still jaguars in the area? Yes (__) No (__) Don’ t know (__) Where about? ______

2.3 Do you consider the jaguar a problematic animals? Yes (__) Could be (__) No (__) Don’ t know (__) Why?______

2.4 Do you think that is important to conserve jaguars in these lands? Yes (__) Don’ t care (__) No (__) Don’ t know (__) Why?______

2.5 Do you like to share this area with jaguars? Yes (__) Don’t care (__) No (__) Don’t know (__)

2.6 Would you be interested in tourists visiting the area to see wildlife? Yes (__) Don’t care (__) No (__)

2.7 Do you think that jaguars can attack humans without being provoked? Yes (__) No (__) Could be (__) Don’t know (__)

2.8 Have you ever heard of such an attack? Yes (__) No (__) If yes, ______

2.9 Have you seen a jaguar? Yes (__) No (__) Dead/captured (__) When was the last time that you saw one? ______Hour: ______Where? ______

3. Killing of jaguars

3.1 Are jaguars still killed in the area? Yes (__) No (__) Don’t know (__)

3.2 If yes, what are the most common reasons? Conflict with livestock (__) Fear (__) Interest in the skin (__) Sport and entertainment (__) Tradition (__) Conflict with hunters (__) Other (______)

What methods are used? ______

3.3 Over the last year, were any jaguars killed in this area? Yes (__) No (__) Do not know (__) If YES how many? _____

3.4 Could you give me an estimate of how many jaguars were killed in this area in the past? ______

3.5 Could you tell me about jaguar hunting in areas sorrounding this one? ______

3.6 Do you think jaguars are more, less or the same than before? More (__) Less (__) Same (__) Do not know (__) 3.7 Why? ______

Appendix III

III.I Photo-inventory of mammal and bird species recorded in the study area (The bat and mouse were not included because they were not identified to the species level)