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PHILIPPINE EAGLE Pithecophaga jefferyi BREEDING BIOLOGY, DIET, BEHAVIOR, NEST CHARACTERISTICS, AND LONGEVITY ESTIMATE IN MINDANAO ISLAND

A Thesis Presented to the Faculty of the Graduate School Ateneo de Davao University Davao City

In Partial Fulfillment of the Requirements for the Degree Masters of Science in Biology (Zoology)

Jayson C. Ibaez March 2007

APPROVAL SHEET

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ACKNOWLEDGMENTS

Without the assistance and cooperation of my colleagues at the Philippine Eagle Foundation, I would not have completed my thesis. I am grateful to my research team namely Mary Beal Jiloca, Camille Concepcion, Aniceto Allado, Adriano Oxales, Giovanie Tampos, Maita Verdote, Medel Silvosa, Jo Cruz, Guiller Opiso, Joseph Alcomendras and Perfecto Balicao. During the 1999 fieldwork, Donald Afan, Glen Bueser and Kharina Bueser helped in data gathering. They were all very consistent and passionate in observing the eagles in the wild. Dennis Salvador, my organizations Executive Director, has been very supportive and patient with me. Dr. Hector Miranda, former PEF Science Director, colleague and a friend has been very influential in my career and has inspired me to start observing breeding Philippine Eagles in 1999. I also thank my co-managers, Domingo Tadena and Lito Cereno who were my mentors and inspiration for my work with the Philippine Eagles. I am grateful and very indebted to Prof. Fe Bagajo, my Thesis Adviser. She has been an exceptional academic mentor whose continued support, inspiration, attention and advising have been instrumental to my success at Ateneo de Davao University. I am forever thankful to my colleague, friend and my wife, Bing Ibanez as she was my field assistant during the 1999 field work, and more importantly has provided support, friendship and inspiration. Dr. Nigel Collar was very generous to host my internship at Cambridge University. He has provided intellectual stimulation and a refuge that provided the right academic atmosphere for thesis writing. This thesis would not have been written without his guidance. My research committee members: Dr.

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Bernadette del Rosario, Dr. Cecile Clarinda Pasino, and Dr. J-ney Zapanta. They all made important academic guidance and contributions. I also thank Dr. Todd Katzner for the important comments over the manuscript. I thank the local government units, Indigenous Peoples, and friends at Arakan, Mati, Tarragona, Sibulan and Lantapan for providing permission and assistance to do research within their locality. Before I worked in Davao City, my undergraduate mentor Prof. Andy Dans generously embraced me into his team and opened opportunities for work and further learning for me. I am forever grateful to him and his family. My enrollment to the Masters Degree was supported by the Marubeni Energy Services and the Philippine Eagle Foundation. Field expenses were supported by the Marubeni Energy Services, the Peregrine Fund, the Critical Ecosystem Partnership Fund, and the Philippine Eagle Foundation. The Department of Zoology, Cambridge University financed my internship at Cambridge UK. I also acknowledge the assistance of undergraduate students Mayette Sulapas and Allen Conte. Thanks also to our field trustees who watched over the eagles and their nests whenever researchers were away: Romeo Abe, Blackie Lomiston, Melanio Dindin, Manolo Labonso, and Nene Balicao. Support and inspiration were also provided by my family and friends and by many colleagues in the conservation movement. To all of those people and organizations that Ive inadvertently omitted, thank you. Finally, this thesis is for happiness, which in my case would mean Bing, Sofia and Luis.

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ABSTRACT

This study described the breeding biology, diet, behavior and nest characteristics of Philippine Eagles in Mindanao using data from five Philippine Eagle pairs nesting from 1999 to 2007. Using information on breeding success spanning three decades (1977-2007), an estimate of longevity for Philippine Eagles was also calculated. Although results for breeding behavior, diet analysis, and nests and nest tree characterization did not vary considerably from previous studies, this study provided additional details and insights from the parameters considered. Data showed that the Philippine Eagle has the longest and energetically most expensive parental investment for any birds of prey. Baseline values for nest attendance, incubation and brooding bout duration, trip duration and feeding rates during each stage of breeding for each sex were provided. Nest trees were located near human communities and were at varying distances from the forest edge. Measurements of nests and nest trees were not very different from previous nests, although surface area of nests studied were larger on the average. Flying lemur Cynocephalus volans was the primary prey species, and arboreal mammals was the most important prey group. Because the populations of Mindanao arboreal mammals seem stable, these animals will most likely remain as the important prey group on the island. At least 14 prey taxa were recorded and 9 were identified to the species level. Philippine Eagles are opportunistic feeders that can shift their diet to what is available. When heavier, native mammals are scarce, they seem to adjust by taking on smaller prey items at a higher frequency. An explanation for sexual dimorphism in raptors predicts that female

take larger and heftier prey whereas the males on smaller, more agile prey. The study found no evidence for prey base partitioning among sexes in Philippine Eagles. This study documented the first evidence of predation on domestic animals and pets, and two instances of breeding failures because of food stress. In order for adults to produce sexually mature birds which will replace them when they die, each male and female must live at least 29 to 30 years. The fact that the Philippine Eagle is long-lived, invests a lot on breeding, and the continuing habitat loss and persecution predict the Philippine Eagles vulnerability to extinction. The species also nest along forest edges near upland communities and can feed on domestic animals and pets. These make the eagles very susceptible to shooting, trapping and other forms of human persecution. An egg and a nestling were deserted most likely because of food stress. However, there are evidences that eagle pairs in large, intact forests with stable numbers of native arboreal mammals are breeding well. Because of the importance of increasing adult survival as well as ensuring productivity, protection of breeding adults and the places where they nest are important. Conservation of the home range where they hunt is equally important to ensure that enough prey base is available. Community-based conservation (CBC) approaches can be a potent tool for conservation in places where people and Philippine Eagle conflicts are tense.

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TABLE OF CONTENTS Content TITLE PAGE APPROVAL SHEET ACKNOWLEDGEMENT ABSTRACT TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES CHAPTER 1 INTRODUCTION Background of the Study Statement of the Problem Significance of the Study Scope and Delimitation of the Study Definition of Terms REVIEW OF RELATED LITERATURE AND STUDIES Value of Conserving the Philippine Eagle Conservation Status of the Philippine Hatching and Fledging Success Mortality Nesting Density and Population Estimate for Mindanao Entire Philippine Population Estimate and the Luzon Dilemma Breeding Biology and Ecology of Philippine Eagles Breeding: Monogamy and Site Fidelity Breeding: Eggs, Timing of Breeding, and Incubation 11 11 12 13 14 16 17 17 19 1 4 5 8 9 Page No. i ii iii v vii xi xii

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Nesting Behavior: Courtship and Pre-Egg Laying Nesting Behavior: incubation and brooding Growth and development of young Diet and Feeding Regime Predation on Domestic Animals Nest Sites and Nest Trees

20 21 22 24 25 27

METHODOLOGY Description of Study Area Field Data Collection Data Analysis PRESENTATION, ANALYSIS AND INTERPRETATION OF DATA Nesting Phenology Nest and Nest Tree Characteristics Reproductive Success Behavior of Adults Pre egg-laying period Courtship displays Incubation stage Sprig deliveries Behavior on the nest Prey delivery rate Brooding stage Sprig deliveries Behavior on the nest Prey delivery rate Post-brooding stage Sprig deliveries 38 39 43 44 44 44 47 47 48 50 51 51 51 53 54 54 29 33 35

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Adult behavior on the nest Prey delivery rate Diel pattern of prey delivery Parental care Nestling Development and Behavior Diet and Food Habits Prey species and food habits Philippine Eagle hunting techniques Predation on domestic animals Philippine Eagle diet and food habits at Apo and Kitanglad nests Size difference between prey items hunted by male and female eagles Breeding failures as an effect of food stress Estimates for Philippine Eagle adult longevity

57 57 61 63 64 66 66 70 71 73 75 77 78

SUMMARY, CONCLUSION, AND RECOMMENDATION Summary Conclusion Recommendations 79 81 82 84 90 95 96 99 100 101

REFERENCES APPENDICES A B C D E F Photos of Philippine Eagles, nests and/or nesting sites. Data sheet used in studying Philippine Eagle behavior on the nest. Philippine Eagle Ethogram. Sample of transmitter attachment and window clipping method to mark Philippine Eagles Mean weight used for calculating biomass of prey items and the references. Outcomes of breeding attempts by Philippine Eagle pairs in Mindanao from 1977-2007.

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G H I

Calculation of longevity value for Philippine Eagles in the wild state in Mindanao. 2008 weights of male and female Philippine Eagles at the Philippine Eagle Center, Davao City. Mean weight of male and female birds in several moderate to highly dimorphic birds of prey.

106 107 108 109

CURRICULUM VITAE

LIST OF TABLES

Number 1 2 3

Title Local land uses within the five Philippine Eagle nesting sites studied from 1999-2007. Nest sites, season and breeding stages studied in Mindanao from 1999-2007. Actual (complete dates) and estimated (month) occurrences of egg laying, hatching and first flight off nest at four Philippine Eagle nests in Mindanao from 1999-2007. Physical characteristics of Philippine Eagle nests and nest trees on Mindanao Island (all measures were in meters). Comparison of sex roles for the incubation stage during daytime observations at three Philippine Eagle nests in Mindanao. Nest attendance, mean trip duration, brooding bout duration and standing bout duration during daytime observations at Sinaka and Kitanglad nests in Mindanao. Prey delivery rates during the brooding and postbrooding (early and late) stages of the nesting period in three Philippine Eagle nests in Mindanao. Quantity of Philippine Eagle prey species observed directly from courtship through the nestling stages at four nest sites in Central and Eastern Mindanao, Philippines, from 1999 to 2007. Food-niche breadth (FNB), dietary overlap, and estimated mean weights (g) of prey of Philippine Eagle pairs at Kitanglad and Mt Apo during the nesting season in Mindanao.

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LIST OF FIGURES

Number 1 2 3

Title Location of five (5) Philippine Eagle nesting sites studied in Mindanao, Philippines. Sprig delivery rate at Kitanglad and Mt Apo nests during the post-brooding period. 18 = 18-27 days, etc. Times of the day sprig was delivered at Kitanglad and Mt Apo nests during the post-brooding period. 5 = 05000559 hr, etc. Comparison of the mean (n=3) for male, female and combined male-female values for the number of prey individuals delivered per day to similar measures for each of three nests (Cabuaya, Apo, Kitanglad) in Mindanao. Comparison of the mean (n=3) for male, female and combined male-female values for average biomass delivered per day to similar measures for each of three nests (Cabuaya, Apo, Kitanglad) in Mindanao. Diel pattern of prey delivery at 3 Philippine Eagle nests during the post-brooding period in Mindanao. 6 = 0600 0759 hrs, etc. Diel pattern of delivery according to weight of prey at 3 Philippine Eagle nests during the post-brooding period in Mindanao. 1 = 1 499 g, etc. Percentage of time spent on diurnal activities by the nestling during the post-brooding behavior at the Cabuaya, Apo and Kitanglad nests in Mindanao. Comparison between the prey mass categories brought by male and female during the post-breeding phase in three Philippine Eagle nests observed (Cabuaya, Mt Apo, Kitanglad) in Mindanao. 1= 1499 g, etc.

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Chapter 1

INTRODUCTION

Background of the Study

What is man without the beast? If all the beasts are gone men would die from great loneliness of spirit. For whatever happens to the beasts also happens to man. All things are connected.Chief Seattle (Duwarnish Tribe). If there is one statement that would explain the value of conserving threatened species, this wisdom from an Indian Chief will suffice: people must conserve wild animals for the benefit of humankind. But a lot of our unique wildlife is facing extinction, not by natural causes but by the irresponsible human exploitation of their kind and the natural places where they live. These creatures at the brink of extinction include our national bird - the Great Philippine Eagle. Called the Haring Ibon (English King of Birds), The Philippine Eagle (Pithecophaga jefferyi) is a tall, huge, broad-winged tropical forest raptor endemic to Luzon, Leyte, Samar and Mindanao islands of the Philippine archipelago. As a long-lived, late maturing, slow breeder heavily persecuted in a forest habitat that is rapidly disappearing, the Philippine Eagle is one of the worlds rarest and most highly threatened bird species (Birdlife International, 2007). Birds of this species nest on large piles of sticks on epiphytes atop the crown of big trees in a range of habitats from primary to residual forests near human dwellings (Kennedy, 1985). Lowland dipterocarp forest is believed to be

the previous stronghold (Rabor, 1969; Kennedy, 1981; 1985), but recent distribution records show that they occur in mountain forests as well (Collar et al., 1999). The Philippine Eagle is the only species in its genus (i.e. monotypic, Brown and Amadon, 1968), but owing to its massive size, ecology and morphology, it was placed under the harpy eagle group (tribe Harpiinae) together with the eagles of New Guinea Harpyopsis and South American Harpia and Morphnus (Brown and Amadon, 1968). However, recent phylogenetic studies showed that Philippine Eagles are closely related to snake eagles (Circatinae) than to the three forest eagles mentioned earlier (Lerner and Mindel, 2005). Conservation of the species was first brought to world attention in 1965 during a conference by the International Union for the Conservation of Nature and Natural Resources (IUCN) in Bangkok, Thailand (Rabor, 1965). The late Prof. Dioscoro Rabor alerted his foreign colleagues about the impending extinction of the Philippine eagle, with only less than 100 individuals suspected in the wild. Through the World Wildlife Fund, the Philippine government declared the bird as protected a year later. Subsequently, a conservation program was established. Since then, initiatives by the government and non-government organizations to monitor the status and breeding success of known eagle pairs, protect its remaining forest habitats, do conservation breeding of the species, and implement public education programs grew in earnest. Despite the growing public attention on the species, published information on the biology and ecology of the Philippine Eagles remained few. The reasons include fund limitations, the lack of investigators trained in research and scientific

writing, and the difficulty of doing research in often rugged and rebel-infested areas (Salvador and Ibaez, 2006). For example, since the last publication by Dr. Robert S. Kennedy on the conservation status of the Philippine Eagle more than two decades ago (Kennedy, 1981; 1985), only seven papers (3.5 papers in 10 years) were published in scientific journals. All of these were from Mindanao. In contrast, no scientific paper has ever been published on eagle populations since 1985 on the islands of Luzon, Leyte and Samar. Because of this, the Philippine eagle remains one of the least understood large raptors in the world. As a critically endangered species that is facing a 50 % chance of getting extinct in the next 30 years, the Great Philippine Eagle is a global priority for conservation (Bildstein et al., 1998). But in order to save a species, its biological and ecological requirements must first be understood - at least its basic survival requirements so that its well-being can be secured (Burnham and Cade, 1995). It is difficult to save an animal that one does not understand well. In 1999, the Philippine Eagle Foundation (PEF), a non-governmental organization dedicated to the conservation of the Philippine Eagle and its rainforest habitat embarked on a long-term study of the behavior and food habits of breeding Philippine Eagles in Mindanao to address information gaps on the ecology and behavior of the species. This thesis described the results for five breeding pairs studied from 1999 to 2007.

Statement of the Problem The breeding biology, diet and nest characteristics of Philippine Eagles in Mindanao were partially described by Gonzales (1968) and Kennedy (1985). The courtship and pre-egg laying period are yet to be documented. The activity regimes of nesting pairs, including male and female patterns, were insufficiently quantified. Prey species have been identified (Gonzales, 1968; Kennedy, 1985), but only from a limited number of nests. There are unconfirmed local reports of eagle predation on domestic animals. In the light of shooting and trapping of Philippine Eagles as a result of perceptions that they are livelihood pests, measuring the importance of domestic prey in the eagles diet is important. Additional information on nests and nest trees must also be collected as forests have shrunk for the past two decades.

General Objective

This study aims to describe the breeding biology, diet, behavior, nest characteristics and average longevity of the Philippine Eagle Pithecophaga jefferyi in Mindanao Island using data from five nests observed from 1999 to 2007 and from historical data.

Specific Objectives Specifically, it aims to: 1. Describe the nesting phenology of Philippine Eagles breeding in Mindanao, 2. Describe nest and nest tree characteristics, 3. Describe reproductive success, 4. Describe the behavior of adults during pre-egg laying, incubation, brooding and post-brooding stages of the nesting cycle, 5. Describe nestling development and behavior, 6. Describe the diet and food habits of breeding adults and nestling, and 7. Estimate the longevity and productivity of breeding adults.

Significance of the Study

Preventing the extinction of species is a global priority and forms a pillar in the worlds effort to maintain biodiversity (United Nations Environmental Program/Convention on Biological Diversity, 2003). But one can not conserve what one does not know and understand. Basic research on the biology and ecology of threatened species, including knowing its ecological requirements and the factors that limit its population in the wild, is therefore important for protecting small populations and mitigating threats (Burnham and Cade, 1995). This study is a response to a global call for more baseline research on this rare and

extremely threatened species as it was primarily designed to fill in the gap in knowledge on the nesting biology and ecology of Philippine Eagles in Mindanao. This study provided the first detailed study of the nesting biology, breeding behavior, diet and nest characteristics of Philippine Eagles since those by Kennedy (1981,1985) more than two decades ago. Knowing what Philippine Eagles eat is important so prey species can be protected and enough of them exist to meet the Philippine Eagles food requirements. Thus, knowing which animals are the important food for the Philippine Eagles and later using this information to design conservation measures that protect these prey animals and their habitat are the direct benefits of this study. As another aim of the diet study, evidences of domestic animals in the diet of Philippine Eagles was also documented. Many eagles are being shot and trapped by local people because they are seen to be pests of livestock. Measuring the relative importance of domestic animals in the diet of Philippine Eagles is important to properly address the growing conflict between local communities and Philippine Eagles because of alleged eagle predation on livestock. Getting baseline information on how eagles behave during the stages of the nesting cycle, including the time and effort they allot for guarding, feeding and caring for their young is also important to understand how these natural behaviors can be maintained in the face of human impacts on forests habitats. Helping the Philippine Eagles survive in a human-altered world is a big conservation challenge. This study documented the behavior of five pairs only,

which is not enough database to fully understand how Philippine eagles behave in forests under different levels of human influence. But researchers and conservationists can build on the baseline data provided to better understand how eagles behave in response to changes in its forest habitat. This study is supported by the PEF, a non-government organization that works on conserving Philippine Eagle populations. Baseline data will aid PEFs education and habitat protection projects. The results of the study were also used as a basis in developing a species action plan for this endangered bird. The Protected Areas and Wildlife Bureau (PAWB) of the Department of Environment and Natural Resources (DENR) and the PEF convened a national workshop last February 2008 and a 10-year framework of action to save the species from further decline will be published soon. It has been the governments policy to involve stakeholders in the conservation of species and habitats. A number of local government units in eastern Mindanao have started delineating municipal forests as protected areas within their jurisdiction using boundaries of Philippine Eagle territories. Local nongovernmental organizations are also using the Philippine Eagle as a flagship for forest conservation and sustainable development in the uplands. All the information generated will be useful for building better awareness and appreciation of biodiversity among students and will hopefully encourage them to take care of Philippine Eagles and other wildlife for sustainable development.

Scope and Delimitation of the Study The breeding biology, diet, behavior and nest characteristics of Philippine Eagles in Mindanao island was described and interpreted using data gathered from five Philippine Eagle breeding pairs and nesting territories in Central and Eastern Mindanao. These eagle pairs and their nests were studied from July 1999 to July 2007. Five pairs is not ideal. But with the rarity of the Philippine Eagle, the fact that they occur in low densities in often rugged and inaccessible terrain, the costs associated with doing behavioral studies, and the strict government guidelines on studying critically endangered species in their natural habitats, five pairs is a good number. Data on breeding behavior comprised observations on activities during the following stages: pre-egg laying or courtship stage, incubation, brooding, and post-brooding. However, not all of these stages were observed for all of the pairs. The nests were found at different stages of the nesting cycle (e.g. the Cabuaya and Apo nests were found during the post-brooding period) and had different breeding outcomes (e.g. nesting at Tarragona and Sinaka failed during the incubation and the brooding stages, respectively). The post-fledging stage, the last phase of the nesting cycle, will be excluded as there were not enough resources to extend observation beyond the fledging stage. Data on diet was based on the direct observation of prey items brought by breeding adults to the nest. The longevity estimate used breeding success data for Mindanao alone where substantial information from three decades of Philippine Eagle monitoring by the PEF has been compiled.

Definition of Terms Breeding Biology. Refers to the timing and the nature of the various stages in

the breeding cycle of a species. Among birds, these stages would be the pre-egg laying, egg, incubation, hatching, nestling, post-brooding, and the post-fledging periods. Conservation Status. Refers to the category of the species under

consideration according to the International Union for the Conservation of Nature and Natural Resources (IUCN) classification and the efforts done to conserve the species and its habitat Critically Endangered. A species is Critically Endangered when it has a 50 %

chance of getting extinct in the next 30 years, and thus facing an extremely high risk of extinction in the wild. Breeding success. fledging stage. Brooding stage. That stage in the breeding cycle of birds from hatching until Breeding is successful when the young bird reached post-

that period when the parents no longer sits on the young bird to regulate its body temperature. Congeners. Diet. Belonging to the same genus.

Refers to the species of prey consumed by breeding Philippine Eagles. The closest distance between active nests of two

Inter-nest distance.

monogamous eagle pairs. Longevity. The average age by which an adult Philippine Eagle successfully

rears a young that reached sexual maturity.

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Nesting phenology. Nest characteristics. parameters. Nest site fidelity.

Study of the timing of breeding and its various phases. Refers to measures of eight nest and nest tree

Refers to a trait of some species of birds in which they use

the same place for nesting and rearing young several times. Philippine Eagle. A large species of a forest eagle that is found only in the

islands of Luzon, Leyte, Samar and Mindanao of the Philippine archipelago. Post-brooding stage. That stage wherein young birds on the nest can regulate

its own body temperature and thus, no longer depends on the warmth of its parents body. Post-fledging stage. That stage from the time the young bird flew off the nest

until the young disperses away from its parents territory and becomes totally independent. Pre-egg laying or courtship stage. That stage in the breeding cycle of birds

prior to egg-laying which broadly include nest building, courtship and the mating activities.

Chapter 2

REVIEW OF RELATED LITERATURE AND STUDIES

Value of Conserving the Philippine Eagle

Predators, like the Philippine Eagle, are considered good ecological indicators because they are atop food chains and require wide home ranges (Rodriguez-Estrella et al., 1998). Predatory birds or raptors are considered good indicators of habitat quality because of their sensitivity to human disturbance and environmental contaminants (Newton, 1979). In some cases, raptors are used as an umbrella species for biodiversity conservation because their protection results to the protection of all plant and animal species found within their large home ranges (Burnham and Cade, 1995). More importantly, the Philippine Eagle is a national patrimony that must be cared for. It is an apt symbol of the Filipino people, being found only in the country and being the countrys national bird.

Conservation Status of the Philippine Eagle

The conservation and management of any species is contingent upon an understanding of its population dynamics that is, how population number changes through time as reflected in net death and birth rates (McGahan, 1968). Fortunately, breeding records of several pairs in Mindanao from 1977 up to now

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were amassed, providing insights on the success of eagle breeding in the wild, at least in the Mindanao part of its range. For populations in Luzon, Leyte and Samar where no nest or breeding has ever been studied, nothing is known about breeding success rates there. Similarly, death rates are difficult to estimate as the number of eagles dying in the wild remains undocumented. This is true in all of the islands where it is found. But in Mindanao where several wild eagles were turned over to and rehabilitated at the Philippine Eagle Center in Davao City, a trend in the contribution of people to eagle mortalities can be inferred.

Hatching and Fledging Success

From 1978 to 2007, 81 nesting by 30 pairs hatched 73 young for an average of 2.7 young hatched/year. Of those hatched 53 fledged, with an average of 1.8 fledglings/ year. Overall, the breeding success in Mindanao from 1977 to 2006 appears to be 65 % or 0.32 young/ pair/ yr. In a previous summary of breeding success of Philippine Eagles that excluded data from 1999 to 2007, Mindanao Philippine Eagles registered an over-all success rate of 58 % or 0.29 young/ pair/ yr (Miranda et al., 2000). The average breeding success for large eagles is less than one young/pair/year (Newton, 1979). This indicates that Philippine eagles in Mindanao seem to be breeding within normal levels. Thus, it seems that abnormal mortality rates among sub-adults and adults, and not breeding failure, is the proximate cause of decline (Miranda et al., 2000).

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Mortality The only source of information on deaths among wild Philippine Eagles were from the outcomes of rescues and rehabilitation of birds captured by people (Hinlo et al., in press). Out of 59 Philippine eagles that were admitted at the Philippine Eagle Center (PEC) from 1970 to 2006, 36 birds died, 5 were released back to the forest, while the rest were kept as breeding stock. Of the 36 birds that have died, 16 (44 %) had an undetermined cause of death, 9 (25 %) died of infectious disease, 6 (17 %) of metabolic and nutritional disease, 4 (11 %) died of trauma, while one eagle died of cancer. Although the primary causes of most of the deaths among eagles admitted to the center are unknown, these cases are definitely human-caused, directly or indirectly. Shooting and beating eagles that were trapped or cornered in the forest are clearly human-inflicted, but so too are deaths resulting from prolonged stress brought about by improper handling and feeding of trapped or caged eagles by inexperienced captors. Looking at the timing of the deaths in 36 birds, nearly half (16 birds) occurred within a month after admission. In all of these birds, they were either too injured or mishandled to be saved, or too sick and malnourished to be cured (Hinlo et al., in press). Based on this figure, it seems that the minimum mortality rate due to direct human interference is one eagle every year. But if we include the 19 eagles that are alive and in captivity as mortalities as they are already removed from the wild, it seems Filipinos are responsible for losing at least 3 birds in 2 years (1.5 eagles/ yr). But this is admittedly an underestimate as there are surely more

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unreported cases of human-caused deaths, especially in remote areas of Mindanao where news of eagles being shot or dying in captivity do not reach public knowledge. Eagles can also be lost to natural causes such as old age and diseases, but it is unknown how many eagles are dying from these causes. Of the infectious diseases known to inflict eagles, Aspergillosis could be the most important. Out of the 15 eagles whose causes of death at the PEC are uncertain, 10 were unconfirmed Aspergillosis cases. Caused by the fungus Aspergillus fumigates, Aspergillosis is the most common cause of death in captive raptors in Britain and the United States (Cooper, 1985). Golden eagles, gyrfalcons, ospreys, goshawks and the rough-legged hawk are at high risk for this infection (Joseph, 1998). And perhaps Philippine eagles too, as Aspergillosis were responsible for many raptor deaths at the Center for Philippine Raptors in Luzon (Celis R., pers. comm.). A. fumigates are ubiquitous, naturally occurring, and often manifests itself as a sequel to some other forms of stress such as recent change in management, poor condition and concurrent disease/injury (Redig, 1993).

Nesting density and population estimate for Mindanao

Current knowledge on the breeding density and estimate of population in Mindanao is from the work by Bueser and colleagues (2001). By mapping the nests of Philippine Eagles from 8 provinces in Mindanao from 1991 to 1998, the density of adult breeding pairs was estimated at one pair every 127 - 133 km2.

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Mean inter-nest distances was calculated at 12.7 km, while the average distance between breeding areas (i.e. including pairs whose nests were not located ) was 12.95 km. Using the nest space average, Bueser and colleagues (2001) calculated the amount of forest for each territorial space centered on nests using 1:250,000 land satellite interpretation maps (NAMRIA). The average extent of forest cover for each nest area was estimated at 55 percent, or an average absolute value of about 70 km2. The total forest cover for each nesting "space" was approximated to include closed canopy (with more than 50% cover) forest, open canopy (with less than 50 % cover) forest, montane and mossy forest. Old-growth forest averaged 51.0 % for pairs with located nests (N = 13) and 56.0 % for all pairs (N = 25), which includes those whose nests were not found. To determine a population estimate, two forest cover averages were used: a) 68 km2 for pairs with located nests only, and b) 72 km2 for all breeding pairs. Using these densities and the estimated suitable forest area of approximately 13,898 km2, Bueser and colleagues (2001) estimated that there are about 201 19 and 190 23 occupied territories in the island. This estimate is conservative since they excluded sub-adults and floaters" which they said are difficult to incorporate into the data set in the absence of information on survival rates and dispersal. Implicit to this estimate is the assumption that all suitable areas are occupied.

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Entire Philippine population estimate and the Luzon dilemma Similar extrapolations on the population estimate to include the islands of Luzon, Samar and Leyte suggest a total population of 340 pairs (Miranda, et al. 2001). However, the estimate for Luzon may have a large margin of error. Extensive surveys during the late 1970s and early 1980s did not result in locating an active nest (Kennedy, 1985). Although, 81 nestings by 30 pairs have been monitored in Mindanao during the last two decades, no nest has been studied in Luzon, despite the fact that the island has more forest cover than Mindanao. At present, it is arguable whether this is a reflection of differential nesting densities, forest type variation, biogeographical history of the population, or differences in research and conservation efforts (Miranda et al., 2001). Studies on Luzon populations are imperative for many reasons. Luzon may have been separated from the Greater Mindanao (which include Samar, Leyte, and Mindanao) by 10 million years (Heaney and Regalado, 1998). From the standpoint of conservation genetics, a vicariant historical event could have led to genetic differentiation between the Luzon and the Greater Mindanao (Samar, Leyte, and Mindanao) populations. This will be tested in the near future using molecular tools and should be an important consideration in management decisions involving releases of captive bred or rehabilitated birds from one island to another (Miranda, et al, 2001).

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Breeding Biology and Ecology of Philippine Eagles Studies on the breeding biology, behavior and diet of Philippine Eagles are too few. In fact, the foundations of our current knowledge on the nesting biology of the species still rely on only two studies - that by Gonzales (1968) who observed a nesting pair and its young in Malalag, Davao del Sur from July 1963 to June 1964 and those by Kennedy (1981, 1985) who documented 5 nests in different parts of Mindanao from December, 1977 to November, 1979. Data for these are effectively summarized in the book Threatened Birds of the Philippines by Collar and colleagues (Collar et al., 1999). Published notes and accounts of previous investigators and results of similar studies on raptors outside the country were also used to enrich the review below.

Breeding: Monogamy and Site Fidelity

It has been assumed that Philippine Eagles breed as monogamous pairs (Gonzales, 1968; Kennedy, 1977; Kennedy, 1981). Incidentally, this is the most common mating system among raptors (Newton, 1979). Monogamy is believed to have probably evolved among birds to meet the parental care needs of the young, among others (Newton, 1979; Mock, 1985). Philippine Eagles are also believed to mate for life as long as the mate is alive (Kennedy, 1981), but this has never been verified in the wild. No eagle pairs have ever been marked, which is unfortunate because monitoring marked birds has a lot of benefits, including tracking faithfulness to the same mate and

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instances of mate replacement or mate turn-over rates (Young and Kochert, 1987). But eagles in captivity exhibit high mate fidelity. The longest time that a captive eagle pair stayed together in one cage is at least 9 years (Tadena pers. comm), and this pair still lives together. Meanwhile, some Bald eagle (Haliaeetus leucocephalus) pairs which are about two thirds the size of the Philippine Eagle can stay together in the same territory for 6 breeding seasons (Jenkins and Jackman, 1993). For the Eurasian sparrowhawk Accipiter nixus which is about one thirds the size of the Philippine Eagle, four years was the longest that a pair was found on the same territory (Newton, 1979). Harvey et al. (1979) suggested that long-lived birds and birds living in stable environments (e.g. tropical forests) may be more faithful to mates than other birds. There is no question that Philippine Eagles exhibit high fidelity to nesting sites. For example, at Mount Apo in Mindanao, breeding has continued in one core area for more than three decades now. The earliest record was in 1972 (Kennedy, 1977) and the latest during the 2005 season where a juvenile successfully flew off the nest in July 2006 (Concepcion et al, in press). In Mount Kitanglad, the first record of breeding was in 1986 (Miranda et al., 2000) and a pair is still breeding up to now. There are 8 more pairs in Mindanao that also nested at least twice in the same places (Miranda et al., 2001). Nest site fidelity has also been observed in species that also exhibit mate fidelity such as Bald eagles (Jenkins and Jackman, 1993) and in many other raptors such as the Peregrine falcons, African eagles, Great Kestrels, Buteo species, and European sparrowhawks (Newton, 1979). Nest sites that are used repeatedly are found to

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be those places where breeding success is high so that re-use is advantageous for maximizing raptor productivity (Rowley, 1983).

Breeding: eggs, timing of breeding, and incubation

Philippine Eagles lay only a single egg every other year (Kennedy, 1981). Although early records say they can also lay two eggs (Grossman and Hamlet, 1964), it was only in two instances that this happened (Wylie, 1974). A captive bird in San Diego Zoological Garden produced two eggs in 1974, one on April 4 and the other on April 5. The other instance was recorded in 1973 at the Philadelphia Zoological Garden where one was laid on February 3 and the other on February 17. Based on the monitoring of 28 breeding pairs in Mindanao between 1978 and 1998, only a single egg was documented in each nesting attempt (Miranda et al., 2001) which indicates that the previous records are more of a rare exception than a rule. However, captive propagation of eagles confirmed that eagles can indeed lay two eggs in one season, but only when the first one was intentionally removed from the nest (Tadena, pers. comm.). Called double-clutching method, this is being employed to increase the productivity of captive birds. There are no recorded instances yet that double clutching occurs in the wild. But if it does happen, it might only be when the egg is lost early in the breeding season where the eagles could re-clutch. Otherwise, they would nest anew the following year. Based on five nests observed between 1977 and 1978, egg-laying in Mindanao seems to occur from September to November (Kennedy, 1985).

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Incubation lasts between 58 to 61 days (Gonzales, 1968; Kennedy, 1981). But in captivity, incubation lasts 57 to 58 days only. Unlike in the wild where attending adults rests in between incubation bouts, eggs in captivity are artificially incubated for 24 hours non-stop (Tadena, pers. comm..).

Nesting Behavior: courtship and pre-egg laying

This stage in the nesting cycle is not adequately documented for Philippine Eagles. There are only two studies on Philippine Eagle nesting biology so that they can be described in detail and individually. Gonzales (1968) who studied a breeding pair at Davao del Sur in 1973 briefly mentioned that the pair made frequent daily visits to the nest and called more often seven days prior to egg-laying. Kennedy (1985) saw more activities from a pair in Amabel, Mount Apo which he observed from September 13 to October 6, 1980. He recorded food exchanges from male to the female, copulations, collections of sprig or branches, and vocalizations, all of which intensified in intensity and frequency during the observation period. Unfortunately, they had to stop the study and leave because of insurgency problems. These observations, however limited, are consistent with the behaviors associated with the courtship period among other raptors. Food exchange from male to female, also called courtship feeding (see Lack, 1940) is typical when egg laying season approaches and thus, is believed to help the female accumulate enough body reserves for breeding (Newton, 1979). However,

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courtship feeding is also suspected to increase the chances of copulation and also might help in strengthening pair bond (Lack, 1940). Besides the mechanical function of consolidating the nest, sprig or branch collection can be a signal to advertise occupancy of a nesting territory, or a form of sex solicitation display as most copulation occurs during bouts of nest building (Newton, 1979). For greeneries, Newton (1979) suggests it could be a form of sanitation or a way to maintain humidity in the nest. On the other hand, Winberger (1984) thinks that nest greeneries may repel nestling ectoparasites. Other courtship displays known for other raptors include elaborate aerial displays, nest displays, greeting and solicitation ceremonies, pre-copulation feeding displays, among others (Newton, 1979; Brown and Amadon, 1968). All of these have yet to be documented for Philippine eagles.

Nesting Behavior: Incubation and Brooding

Philippine Eagles share the time for incubating the egg and caring for the young, but the female spends more time and effort than the male. Gonzales (1968) recorded that the male incubated the egg for less than one third of the total incubation time recorded. However, he did most of the hunting during the nesting. Kennedy (1981, 1985) also had the same results wherein the male did most of the hunting from incubation until the first third of the nestling period, and the female doing about two-thirds of the incubation (apparently also incubation at night) and almost all of the chick-feeding duties during this period. Nest relief ceremonies were also recorded by both investigators and are composed of the

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male delivering food to the female off the nest and immediately taking on the incubation duty while the female eagle feeds and grooms. When the female is ready to return to the nest, she would call. Both studies also showed that the young was brooded, protected and cared for until it was between 40 to 49 days old (Kennedy, 1985; Gonzales, 1968). Thereafter, both the male and the female hunted for food and share in delivering food to, and feeding, the young.

Growth and development of young The young birds studied by Gonzales (1968) and Kennedy (1981, 1985) varied in terms of the periods by which they showed patterns of behaviors. Kennedy first observed the chick doing wing flapping exercises when it was 4 weeks old while Gonzales did not see this happening until the bird is six weeks old. The young birds were observed out of the nest and exploring immediate branches at 15 weeks and 16 weeks by Gonzales and Kennedy, respectively. Gonzales missed describing this but Kennedy first saw the eagle feed by itself when it was 54 days old. The young bird of Gonzales left the nest permanently when it was 15 weeks. One bird observed by Kennedy left the nest when it was 18 weeks old while another when it was 23 weeks old. The post-fledging period is when young eagles learn to fly and hone hunting skills prior to independence and dispersal from its parents territory (Brown and Amadon, 1968; Newton, 1979). Gonzales (1968), Kennedy (1985) and relatively recent observations by Afan and his colleagues (2000) are our best source of information for juvenile behavior during this period.

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Both parents continue to provide food for the young during the postfledging period, but provisioning gradually wanes until the young becomes totally independent. Kennedy (1985) first observed a successful kill of a young eagle when it was 43 weeks old (11 months). Afan and his colleagues (2000) first saw a young eagle grabbed a small unidentified prey from a tree knot hole when it was about 12 months old. For the same bird, it was provided with food by the parents at least 7 times from September 1998 to February 1999 (Afan et al., 2000). The post-fledging stage is also characterized by frequent flight exercises, and other play behaviors. Afan and colleagues (2000) gave a detailed description of a young birds flight repertoire which included gliding, flap-gliding, powered flight, soaring and holding on to branches and vines upside down with wings constantly flapping. The young bird was also seen taking branches and sticks and playing with it as if it was a prey item being subdued or a material being arranged on an imaginary nest bowl. All of these were interpreted as play activities that improve flight techniques and hone hunting skills. Such play activities happened in the absence of the parents, suggesting that hunting techniques appear to be learned by the young independently. Young eagles

were mobbed by birds such as large-billed crows Corvus macrorynchus (Gonzales, 1968; Afan et al., 2000) rufous hornbills Buceros hydrocorax (Gonzales, 1968; Kennedy, 1985) and writhed hornbills (Afan et al., 2000), but managed to stay safe sometimes with help from the parents (Gonzales, 1968).

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The precise age the young becomes independent from its parents is yet unknown as there has been no detailed and continuous study of the whole postfledging period in Philippine Eagles. Kennedy (1985), however, observed that a day after a young eagle was last seen within the territory, the parent eagles started copulating. They last saw the young eagle when it was 16 months and 22 days.. Whether or not the parents actively drive young eagles out of the territory remains an open question. But in some large eagles like the crowned eagle in east Africa, the young seem to leave on their own accord (Newton, 1979).

Diet and feeding regime

Our best source of information on the diet of Philippine eagles at least during the breeding season is Kennedy (1981, 1985) with his study of the prey species brought by 5 adult pairs to the nest in Mindanao. Fourteen vertebrate prey items were recorded with flying lemur (Cynocephalus volans) as the most frequent prey species delivered to the nest (54.2 % of identifiable prey). Palm civet (Paradoxurus hermaphrodites) and Mindanao flying squirrel (Petinomys crinitus) were the next most abundant prey items at a frequency of 12.0 % and 7.8 %, respectively. The rest were 3 species of birds, at least 3 species of snakes and 2 species of lizards. The top five prey items brought to the nest according to live weight were the young Philippine deer (Cervus mariannus) (13-14 kg), the long-tailed macaque (Macaca fascicularis) (3-4 kg), palm civet (3-4 kg), Rufous hornbill (Buceros hydrocorax) (1.8-2.3 kg) and the Flying Lemur (1-2 kg). In

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contrast to the old generic name as the monkey-eating eagle, the species feeds on monkeys (i.e. long-tailed macaques) infrequently. Philippine eagles have a broad prey base and appear to be opportunistic feeders (Kennedy, 1985), which is similar to at least two equally huge tropical eagles in Africa and in central and south America. Mitani and colleagues (2001) studied 2 pairs of the Crowned hawk-eagles Stephanoaetus coronatus (weight = 3.6-5.0 kg) in Kibale National Park, Uganda and they identified 16 vertebrate prey species from prey remains, with primates being the primary diet, and bats, rodents, hornbills and unidentified non-primate mammals and birds as minor prey. The live weight of its prey items range between 1.0 to 14 kg. Rettig (1978) studied nesting Harpy eagles Harpya hapija (7.5-9 kg) in Guyana and found the same prey size variety. Harpy eagles fed on 15 vertebrate taxa and took animals with live weights between 2.7 to 9.0 kg. All of the prey items though were mammals. All these giant eagles share a preference for arboreal mammals (i.e. >60 % of prey): flying lemurs for the Philippine eagle, primates for the Crowned hawk-eagles and sloths and primates for Harpy eagles. All also exhibit the same ability to prey on animals much heavier than themselves.

Predation on domestic animals

Local reports that Philippine Eagles take domestic animals as prey are common in almost every place where they are found. They allegedly take domestic pigs, young goats, puppies, cats and chickens, especially during the nesting season. But this has never been verified. The earliest record of local

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accounts is those by John Whitehead in 1896 who wrote in his field notes that it is well known to the natives as a robber of their poultry and small pigs (Whitehead, 1899). However, in previous observations of several nests in Mindanao, no domestic animals were ever brought to the nest by the attending adults (Gonzales, 1968; Kennedy, 1981; 1985). Raptor predation on domestic animals is well documented and is the subject of mitigating actions by local governments in US and Europe. For example, Golden eagles (Aquila chrysaetos) allegedly fed on lambs (Ovis aries) and a calf (Tigner and Larson, 1981; Wood, 1946), Bald eagles (Haliaeetus leucocephalus) on sheep (McEneaney and Jenkins, 1983), the Black vulture on sheep and young pigs (Sus scrofa) (Avery and Cummings, 2007; Lovell, 1947), and the red-tailed hawk Buteo jamaicensis on sheep (Barney, 1959). Because of these, raptors have been seen as nuisance and some resort to shooting and trapping to get rid of them (Kenward, 1999). In aid of management, domestic predation was studied in detail (Phillips and Blom, 1988; Davies, 1999) and recommendations were provided to prevent further losses (Matchett and O'Gara, 1987; Kenward, 1999). Philippine Eagles can take domestic animals as prey. When Philippine eagles were trapped for radio-telemetry from 1998 to 2001 for example, domestic rabbits were used as baits to lure them. Out of 6 eagles that were fitted with transmitters, 4 were trapped using domestic rabbit (Oryctolagus sp.) as bait (Miranda and Ibaez, 2006). Eagles feed on palm civets, wild pig, and a young deer, which closely resemble domestic cats, backyard pigs, and young goats,

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respectively. Captive eagles at the Philippine Eagle Center are fed with meat from goats, chickens, rabbits, Guinea pigs, and quails and they seem to breed successfully with this diet. Philippine eagles, therefore, appears very much

capable of consuming domestic animals so the issue then appears to be not whether they kill domestic animals but to what extent does this occur and under what circumstances.

Nest sites and nest trees

In all recorded instances, from published description and from personal observations, nests are typically located in the crown of huge trees on the major forking at the end of the trunk or along large horizontal branches. Nests consisted of many sticks on huge epiphytic ferns, on large orchids or on a small bed of ferns (Kennedy, 1985). Kennedy measured two nests and each had a surface area of 1.13 x 2.74 m and 1.16 x 1.31 m, respectively. Kennedy also measured two trees and each had a height of 46.0 and 44.7 m, respectively. The nests observed by Kennedy were located in a range of habitats from virgin to residual forests. Formerly known to be dependent only on mature forests (Rabor, 1968), nests found in the last two decades were in a variety of habitats ranging from unlogged old growth forests, to logged forests, a mature agro forest and a forest plantation. In a study of the breeding density of Philippine Eagles in Mindanao, a total of 13 nests were visited (Bueser et al., 2001). And out of this, 3 nests were in primary dipterocarp forest while ten pairs were found on mature secondary

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growth dipterocarp forest. Four pairs nested in the forest interior while nine pairs built their nest less than 100 meters closer to forest edges. Surrounding and within the eagle areas are mosaics of dipterocarp forest at various states of degradation and human-induced modifications characterized by agricultural lands, agroforestry and grassland. Physical parameters of these 13 nests however, were not measured.

Chapter 3

METHODOLOGY

Description of the Study Area Five nests were studied in Mindanao between 1999-2007 breeding seasons as part of the PEFs Philippine eagle monitoring program. Two nests were along the Eastern Mindanao Biodiversity Corridor, two were within protected areas, and one was in a small forest fragment (Fig. 1). Climate in all nests except one falls under a Type IV rainfall pattern wherein rains are typically distributed evenly throughout the year. Mt Kitanglad has a Type III rainfall pattern wherein November until April is typically dry while the rest of the months are wet. Mt Kitanglad had a temperature range of 10.527C, and an average annual rainfall of 3,800 mm (Heaney et al., 2006). These temperature and rainfall data are probably typical for all mountains in Mindanao. The nest sites are near human dwellings and the nature of land use and extent varied (Table 1). Each nest site is described below (see Appendix A for photos). Mt Sinaka, Arakan, Cotabato Province. Mt Sinaka (7 22.68 N, 125 12.83 E) is an isolated, mid-elevation mountain range (peak at 1448 m) covered by 19 km2 of forests. Only about 5% of its forest is lowland (0-800 m) and most of it is modified by agro-forestry practices. About 85% is montane forest (800-1,200 m) and the rest is mossy forest. Balobo (Diplodiscus paniculatus) and Lamio (Dracontomelon edule) dominate the nesting area, with Malugay (Pometia

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pinnata) and White Lauan (Shorea contorta) making up a minor component. The nest tree was a White Lauan. Patterns of human land use include shifting cultivation for cash crops, maintenance of coffee and fruit trees and, recently, palm oil cultivation. Table 1 Local land uses within the five Philippine Eagle nesting sites studied from 19992007. Nest Sinaka, Arakan Local land use Agroforestry (e.g. shade coffee, fruit tree propagation), slash and burn farming Cabuaya, Mati City Slash-and-burn gathering, hunting Apo, Davao City Slash-and-burn farming, shade coffee, abaca collection, hunting Tubaon, Tarragona Slash-and-burn farming, abaca collection, hunting, rattan gathering Kitanglad, Lantapan Experimental pine plantation, wildlife hunting farming, abaca collection, rattan

Cabuaya, Mati City. The nest (06o 32.63' N " 126o 1.60E) is on a low ridge in a lowland dipterocarp forest facing the east coast of Mt Hamiguitan. Mt Hamiguitan is a mid-elevation mountain range (peak 1,680 m) covering about 319 km2 of dipterocarp, montane, mossy and serpentine forests. Vegetation at the nest site is secondary growth dipterocarp forest, which was subjected to selective logging in the past. Decaying logs and tree stumps provide evidence of these previous logging activities. Large clearings were also present as a result of slash-and-burn agriculture. The forest floor had a thin layer of dry leaf-litter with a humus substrate, and large boulders interspersed the forest floor.

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Mt Apo, Toril, Davao City. The nest (06o 57.85 N, 125o 22.56 E) is located at the eastern border of a strict protection zone within the Mount Apo Natural Park (peak at 2,954 m). Mount Apo is a protected area with a total land area of 721 km2 (Mallari et al 2001). The nest site is about 30 km south-west of Davao City and is being proposed as a 1,430 ha Habitat Management Zone within the park. The active nest, a tanguile (Shorea polysperma,

Dipterocarpaceae) was close to an abandoned farmland right at the forest edge. Approximately 75% of this proposed zone is covered with both secondary and primary dipterocarp forest. Tubaon, Tarragona, Davao Oriental. The nest (N 07 06.40 and E 126 20.87) is confined above the Kamanamayan River, on a steep slope in a mountain range that includes the hills of Piagkiawan, Kasumpo, Painguban, Maningkohon, Pagbawian and Sayapo. The forests of Tarragona are within the Mount Tagub-Kampalili key biodiversity area (KBA) with an estimated forest cover of 1,699 km2. Vegetation is predominantly lowland dipterocarp forests under varying successional stages. Patches of corn and coconut plantations are found close to the forest edges. The nest is within a 1,200-ha Barangay Forest Management Site and a 2,400-ha Community-Based Forest Management Site. Mount Kitanglad, Lantapan, Bukidnon. The nest (N 08o 3.74, E 125o 59.67) is inside the Cinchona Forest Reservation along the southern borders of the Mount Kitanglad Natural Park (total area = 312 km2, Mallari et al., 2001). The nest tree is locally called igem (Dacrycarpus cumingii). Vegetation at the nest site is undisturbed to moderately disturbed lower montane forest, mixed with

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mature exotic pines planted experimentally three decades ago. Terrain ranges from relatively flat to gently sloping, with high rising ridges and cliffs completely absent. The nest is about 1000 m away from a road that cuts through the forest.

Field Data Collection Five Philippine Eagle nesting attempts were studied at different stages of the breeding season (Table 2). Two nests were located through a nest reward scheme of finding eagle nests, while the rest were actively searched using lookand-see and foot survey methods (Bibby et al., 1998). Adult behavior during pre egg-laying and nest-building stage were recorded for the Sinaka nest, during incubation at two nests (Sinaka and Tarragona), during the brooding at two nests (Sinaka and Kitanglad), and during post-brooding at three nests (Cabuaya, Apo and Kitanglad). A total of 2,577 hours of observations were spent. Table 2 Nest sites, season and breeding stages studied in Mindanao from 1999-2007. Nest Season Observed 1999-2000 Breeding Stage Observed Pre-laying and nestbuilding, Incubation, early brooding Cabuaya, Mati City Apo, Davao City 2004-2005 2005-2006 Post-brooding Post-brooding Incubation Incubation, brooding and post-brooding 103 916 74 781 Observation hours 703

Sinaka, Arakan

Tubaon, Tarragona 2006-2007 Kitanglad, Lantapan 2006-2007

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Observations for aerial and other courtship displays were done at Sinaka from open ridges and hills using a 25-60 x field scope and 8 x 40 binoculars. Nests were observed from hides on trees and on the ground at least 60 m away using field scopes. Observation periods were typically from dawn to dusk except in several instances of heavy fog and rain. At the most, two observers occupied each hide and recorded behavioral data at an interval of 10 min (i.e. instantaneous sampling, see Lehner, 1979) using standard data sheets (see sample in Appendix B). A pre-existing list of Philippine Eagle behavioral units (i.e. ethogram developed from observing both captive and wild eagles, see Ethogram in Appendix C) was used. Rare events happening between the 10 min interval were also recorded. Philippine Eagles are sexually dimorphic-females being larger than the males (Brown and Amadon 1968)-so that sexes were distinguished with a high degree of confidence. Apart from the disparity in size, behavioral cues (i.e. female doing most of the incubating, brooding and feeding tasks, male doing mainly hunting during the incubation and brooding stages) helped in differentiating the sexes. During post-brooding observations in which the female helped out with hunting, familiarity with distinct features (e.g. general frame, coloration, size of legs relative to body, etc), behavioral idiosyncrasies (e.g. male at Sinaka relatively uneasy whenever on the nest and near the fledgling), and size of each adult relative to the fledgling complemented the means to distinguish males and females with high certainty. At the Sinaka nest, the female bird had a radio transmitter and a window clipped on her left wing (Appendix D).

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All prey items brought to the nest were identified to the lowest taxonomic category possible. For every item brought, the following data were recorded: the time and weather it was brought, which adult delivered it, body parts that were missing, whether it was fully consumed, and whether prey remains were removed from the nest. All nests and nest trees were measured using eight parameters, with the exception of the Sinaka nest which was measured using two parameters only. Nest height was measured from the level of the rim to the ground. The Nest tree diameter was measured at breast height using a diameter tape. A clinometer was used to measure Nest tree height and a tape measure was used to measure nest width, nest length, nest breadth, nest support branch diameter and nest distance from trunk. The nest was climbed using standard tree climbing and rappelling gear. Geographic locations of nests were taken using hand-held Garmin Global Positioning System (GPS) and mapped using ARCGIS 9.1 (ESRI).

Data Analysis Adult behavior patterns across the nesting phases were described using the following information: 1) daily sprig delivery rates, 2) daily prey delivery rate, 3) nest attendance - percent of instantaneous samples where adult was incubating and brooding, 4) trip duration mean time the egg and the chick was unattended, and 5) incubation and brooding bout duration total time spent incubating or brooding. A bout of standing near the chick during the brooding

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stage was also measured. Diet was described by measuring the percentage frequency of each prey taxa - the no. of individuals in each prey category/total no. of prey in the sample, and the percent prey biomass no. of individuals multiplied by the mean weight of that prey/total prey biomass (see Marti, 1987). Food niche parameters were also compared between two Philippine Eagle pairs. Mean weight of prey (MWP) was estimated to compare average biomass of prey for each pair. MWP was measured by multiplying the total number of individuals for each prey category by the average weight for that category, and then summing these totals and dividing the sum by the total number of prey observed (Marti, 1987). Values for average weight came from either published records for the prey taxa or from actual measurements of sample animals. For prey species for which published weights were absent, weight records of animals close in appearance and size were used (see Appendix E). Food niche breadths (FNB) for each nest were also calculated. FNB was estimated using Levins (1968) equation: FNB = 1/Pij2, where Pi is the proportion of the ith prey category of species j (Marti, 1987). Dietary overlap (i.e. similarity in prey species consumed) was measured between the two pairs using Piankas Index (O): O = PijPik/ Pij 2Pik2 where Pi is the proportion of the ith prey category of species j and k (Marti, 1987). To emphasize the importance of adult mortalities and its implication to Philippine Eagle conservation, an estimate of the longevity and productivity value for Philippine Eagles was attempted using Browns (1966) equation: Longevitye = 2/(N/A [SMy] + age at sexual maturity), where N/A = breeding success (N = no.

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of fledged young, A=no. of attempts reaching egg-laying) and SMy = number of young that reaches sexual maturity (considering 75 % mortality or 25 % survival). The Philippine Eagle breeding information summarized in Miranda et al (2000) was complemented with recent PEF data from 1998 to 2007 to calculate a breeding success estimate for the species (see Appendix F). In the absence of actual data for Philippine Eagles, published values for the percentage of young that reach breeding age were taken from results of ringing Buzzards Buteo buteo and Ospreys Pandion halieatus. The equation was adopted from Brown (1966) who employed the formula for several tropical raptors in Kenya (Appendix G).

Chapter 4

PRESENTATION, ANALYSIS AND INTERPRETATION OF DATA

A. Nesting Phenology

Egg-laying occurred between the months of November to January at four nests (Table 3). It happened on November 1999 at Sinaka, and November 2006 at the Kitanglad nest if one used the 58 days incubation period and counted back from when the egg was hatched (January 21). At Cabuaya, the chicks age was about 4 months old when the nest was found. The young was already full grown then and was no longer brooded by the female. At the Apo nest, fieldwork in January 15-16 and April 4-5, 2006 verified an egg and a three-week old chick, respectively. Using these information, it was surmised that the eggs were laid at the Cabuaya and Apo nests on December 2004 and January 2006, respectively. The Tarragona nest was already with an egg when it was found. But since the egg failed to hatch, it was difficult to estimate when the egg was laid. The egg hatched in January 13 at the Sinaka nest after 58 days of incubation, and in January 21 at the Kitanglad nest (Table 3). Using the 58-day incubation period, hatching could have happened in February and March at Cabuaya and Apo nests, respectively. The nestling period at the Kitanglad nest lasted exactly 144 days (i.e. 4 months and 24 days). The young at Kitanglad, Cabuaya, and Apo fledged in June 2007, July 2005 and August 2006, respectively.

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Table 3 Actual (complete dates) and estimated (month) occurrences of egg laying, hatching and first flight off nest at four Philippine Eagle nests in Mindanao from 1999-2007. Nesting event Egg laid Egg hatched First flight off nest Sinaka Nov 16, 1999 Jan 13, 2000 Cabuaya Dec, 2004 Feb, 2005 Jul, 2005 Apo Jan, 2006 Mar, 2006 Aug 2, 2006 Kitanglad Nov, 2006 Jan 21, 2007 Jun 15, 2007

If the previous record of nesting in September by Kennedy (1985) is included, egg-laying appears to occur within a 5 month window, from September to January. Using data from this study and previous data (Gonzales, 1968; Kennedy, 1985), incubation lasts from 58-61 days. This is so far the longest incubation period among raptors (Newton, 1979). Using previous and recent data, the nestling stage can last between 105 163 days old. Post fledging period according to Kennedy (1985), lasts until the eaglet is 640 days old.

B. Nest and nest tree characteristics

The five nests were at elevations between 630 1,434 m. The Kitanglad nest is the highest in terms of altitude and is in a mid-montane forest that has mature stands of exotic pines. In terms of forest cover, all except one were in forests with size 30,000 km2. The exception was Sinaka with only 19 km2 of forest cover. Four species of trees were used namely White Lauan (Shorea

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contorta, Sinaka), Red Lauan (Shorea negrosensis, Cabuaya ), Tanguile (Shorea polysperma, Apo and Tarragona), and Igem (Dacrycarpus imbricatus, Kitanglad). Nest trees were 3560 m in height (ave. 46.1 13.4 m), and nests were 2046 m from the ground (Table 4). Nest trees were large, with DBH averaging 1.75 m. All nests were on a horizontal branch 28 m away from the trunk. Nests averaged 2.25 m long and 1.4 m wide. Table 4 Physical characteristics of Philippine Eagle nests and nest trees on Mindanao Island (all measurements were in meters, m).
Nest Sinaka Apo Kitanglad Cabuaya Tarragona Average SD Altitude NH DBH TH NW NL NB NSBD NDT

950 630 1434 779 898 938.20 303.30

29.00 0.91 45.92 2.32 38.34 2.34 20.70 1.38 31.67 1.77 33.13 1.75 9.54 0.62

35.40 65.63 43.75 37.81 36.88 43.89 12.56

1.48 1.12 1.90 1.09 1.40 0.38

2.40 2.27 2.76 1.56 2.25 0.50

0.52 0.65 1.20 0.44 0.70 0.34

1.07 0.68 1.80 0.47 1.00 0.58

2.94 1.90 8.13 1.67 3.66 3.03

NH-Nest Height, DBH - Nest tree diameter at breast height, TH-Tree Height, NW-Nest Width, NL-Nest Length, NB-Nest Breadth (Thickness), NSBD-Nest support branch diameter, NDT- Nest distance from trunk.

The five nests were in a range of habitats that were almost similar to the places where Kennedy (1985) and Gonzales (1968) did their observations. The nests were at varying distances from the forest edge, ranging from directly along the edge to about 1,000 m inside the forest. None were in the immediate vicinity of a primary forest. Two were in lowland forests (630779 m altitude), while three in mid-elevation forests (898-1,430 m altitude). All were logged selectively in the

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past and now locally exploited either for agro-forestry, abaca hemp collection, or slash-and-burn farming. Data for nest and nest tree parameters were not very different from measurements made by Gonzales (1968) and Kennedy (1985), but with the exception of measurement for nest surface area. The 1963 nest at Davao del Sur had an estimated dimension of 1.7 1.4 m, while the four nests reported by Kennedy had an average of 1.91 1.25 m. The five nests studied had an average area of 2.25 1.4 m, which is slightly larger than previous nests. But nest heights were within the known range. Gonzales estimate of 28 m and this studys average of 33 m fall well within Kennedys 2746 m range. Nest trees were equally huge with four located in steep slopestwo at the base, one midway and one near a ridgewhile one was in a gently rolling forest terrain. One nest tree species Dacrycarpus cumingi, a non-dipterocarp tree, was previously unrecorded (see summary of nesting tree species in Collar et al., 1999). The eagle pairs studied here and those by Gonzales and Kennedy seem to suggest a preference for forest edges as nesting sites. But such association with edges may be an artifact of the ease of them being found and reported by people. Or, it could be a product of the restricted distribution of suitable nesting trees and substrates for Philippine Eagles. As an effect of massive logging of the lowland forest during the 1960s, most of the large trees are now confined at forest edges in the mountains. Because eagles prefer large trees, they would inevitably end up in forest edges if nesting substrates are limiting.

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Suitable nesting sites are indeed limiting at least in two of the study areas. Largely as a result of logging that peaked during the 1960s, the lowland forests of Apo and Kitanglad have been depleted. What was left were residual dipterocarp forests on steep, inaccessible slopes. In these mountains, eagle preference for forest edges as nest sites seems inevitable. This therefore underscores the importance of managing and conserving actual and potential nest sites at forest edges to ensure eagle productivity. The nest sites were in the vicinity of human settlements, but the Philippine Eagles seem to tolerate human presence. They have repeatedly bred in these sites despite its closeness to villages. The Apo pair seemed to have cohabited with four communities living and farming inside the nesting territory. The earliest record of nesting at Apo was 1977, and eight of the attempts resulted to a fledged young (Miranda et al.,2000, this study). At Sinaka, the adult pair nested six times since 1995 despite its closeness to two communities. These attempts have resulted to four successful fledging (Miranda et al., 2000). The nests at Cabuaya, Tarragona and Kitanglad are 0.51 km from the closest community. Despite human presence in the forests, eagles were constantly detected in Cabuaya during fieldwork that began in 1991 until the nest was finally located in 2005. At Tarragona, the pair re-nested on a new tree close to the 2007 nest at the edge of a slash-and-burn farm. At Kitanglad, the 2007 nesting was the second attempt after the territory was discovered in 2006. Strong fidelity to nesting sites is typical among territorial raptors (Newton, 1979) and appears extreme in Philippine eagles (Miranda et al., 2000; Collar et al., 1999).

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Deforestation trends in Mindanao seemed to have confined several Philippine Eagle nesting sites along forest edges so that several surviving pairs seems to have no choice but to breed near upland communities along these edges. The nesting phase is that part of the eagles life cycle whereby activities of this rather wide-ranging species revolve around caring for a single young. It is within these nest-bound stages of their lives that they become very vulnerable to direct persecution by local people. Because Philippine Eagles exhibit extreme fidelity to nest sites that are close to upland communities, conservation strategies that sustainably meet the needs of the people without compromising the eagles survival and breeding needs as well are therefore critical.

C. Reproductive Success

Three of the nesting attempts by five pairs from 1999 to 2007 resulted to a fledged young, registering a success of 60% or 0.3 young per year. The 1999 attempt at Sinaka failed 9 days into the brooding stage, after three days of intermittent rains and strong winds. The chicks death, apparently because of hypothermia, was confirmed when the female picked the carcass and ate it. For the 2006 attempt at Tarragona, the female wandered away for almost two days and left the egg unguarded. The male delivered only a single prey item to the incubating female within eight days of observation. She left for a long time apparently to feed. Unfortunately, while the female was away, a long-tailed macaque snatched and crushed the egg. This is the first record of a nesting failure caused by a marauding animal in the absence of an attending adult.

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A success rate of 60 % or 0.3 young per year for the five pairs is not very different from a success rate of 58 % or 0.29 young per year for nesting attempts recorded from 1977 to 1998 in Mindanao (Miranda et al., 2000). These figures are not also very different from success rate measured for breeding data from 1977 until 2008 (Appendix F), which is 66 % or .33 young per year. In stable populations of large tropical eagles, an 80 % success or 0.40 young per year is expected (Brown 1966). Based on this data, it seems that Philippine Eagle breeding is not that bad. Thus, our data supports the conclusion of Miranda and colleagues (2000) that breeding failure is not a major cause of decline among Philippine Eagles; at least not yet.

D. Behavior of Adults 1. Pre egg-laying period Courtship Displays. Increased aerial displays and frequent perching near the nest marked the start of the breeding season. From July 7 to September 2 1999, 14 mutual soaring displays over the nesting territory were documented. Such displays were characterized by a soaring flight, circling in the same or opposite directions, with the male usually higher. From great heights, the female performed a diagonal drop with the male trailing in pursuit (dive-chase). It then ended with the male overtaking at great speed, or with the male presenting its talons. In two occasions, the female flipped over and quickly presented its talons as well. Such mutual talon presentation occurs in many large eagles (see Brown

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and Amadon, 1969). The pair also made cruising flights over the territory with the female in the lead. The male also performed advertisement displays, which included calling loudly from prominent perches, and soaring or cruising over the territory (territorial flights). There were male display patterns which included swooping without contact at the perched female (mock attack), calling vigorously in a low, nearly horizontal posture while facing the female (pre-copulation display), and delivering prey for the female to consume (courtship feeding). Courtship feeding happened at a rate of 0.35 prey per day (n=31 days) and was accompanied by copulations. The female did food begs towards the male which were all performed on the nest and were almost identical to sex-solicitation displays by the male. A form of mutual display which may be significant for pair bonding was also recorded. The behavior consisted of repeated pecking at the bill, wings, tail and other body parts of a mate. On one occasion, the male ran its bill along the females wing feathers in a manner similar to preening. Labeled allo-preening, this interaction occurred prior to and after mating, when nest building, or during courtship feeding. The first record of nest material delivery was on September 17 1999, two months before the egg was laid, and it continued all through the pre-laying period at a rate of 1.7 sprigs per day. The male brought more nest materials prior to egg-laying (29 against 22 for the female). Behavior sequence for nest building included (1) acquiring leafy twigs (fresh or dried) or dried sticks (twig without leaves) from nearby trees, (2) depositing on nest, (3) cutting materials into pieces

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and arranging, (4) pressing materials against nest bowl using the breast (breast press), and (5) momentary brooding. The time spent at nest-building prior to egglaying averaged 5.70 4.18 min (n=43). Copulation occurred on nest and nearby perches and ensued with or without courtship feeding. During courtship feeding (or supplemental feeding, see Newton, 1979), the male brought prey to the nest and gave loud and prolonged calls that change into short, excited, and high-pitched whines when the female is close. Calls became vigorous and gained speed as the male mounted the feeding female. Calls waned immediately after copulation. The male mounted the female with its talons curled. If without food, the male exhibits a relatively submissive but wary behavior. With vigorous whining and wings slightly drooping (sex solicitation display), the male walks in a low horizontal posture towards the female. If the female showed no rejecting behavior (i.e. standing erect), the male mounted her. The female eagle remained silent and stooped during copulation. Except for food solicitations, any behavior that suggested sex solicitation on the part of the female was not seen. Copulations were noted on 27 occasions (25 during courtship, 2 during incubation) between September 17 and December 18 1999. It was seen twice in a period of 30 minutes (n=3) and in one instance, it occurred three times in one day. A copulation attempt from mounting till dismounting lasts for 18.83 seconds on the average (SD=2.93, n=6). Copulation among Philippine Eagles seems to be more than what is actually necessary to fertilize an egg. It occurred in a span of three months,

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which is similar to what was observed among Crowned Eagles Stephanoaetus coronatus in Kenya whose breeding biology is similar to the Philippine Eagles (Brown, 1966). Such high copulation frequencies are consistent with reports for other raptors and thought to be more pronounced in raptors than any other bird species (Newton, 1979; Pandolfi et al., 1998; Arroyo, 1999). Several explanations have been proposed. High copulation frequencies strengthen the pair-bond (Newton, 1979). High copulation rates among Bearded Vultures Gepaetus barbatus were thought to buffer the potential risk of extra-pair copulations when the female is away (Bertran and Margalinda, 1999). But such risk is not as important to a monogamous species like the Philippine Eagle. It could be an insurance policy so that when nesting fails early in the incubation, the pair can re-nest immediately. There is no evidence for this yet, but it is known that re-nesting happens at the latter part of the year and not immediately after a failure (Miranda et al., 2000; this study). Copulations also occurred even during the incubation period, from 18 days (Gonzales, 1968) to one month (this study). Continued copulation even after the females fertile period is again thought as a means to strengthen pair-bond (Holt et al., 1992).

2. Incubation stage Sprig deliveries. Delivery of branches to the nests during incubation was exclusively by females at the Tarragona and Kitanglad nests, whereas both adults did it at the Sinaka nest, although the female did most (i.e. 19 of 22 deliveries). Sinaka nest had the highest sprig delivery rate (1.2 per day, n = 19

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days) while nests at Kitanglad and Tarragona, had substantially lower rates: 0.3 per day (n = 6 days) and 0.2 per day (n = 6 days), respectively. Combined, the average delivery rate for the three nests is 0.6 0.6 sprigs per day. Behavior on the nest. In all of the three nests, the female had a substantial share of the daytime incubation duty (Table 5). Out of the total instantaneous samples for each nest, the instances that the female was seen sitting on the egg ranged from 73-83 % (average=82%, n=3). In two nests, however, the male spent more time for each incubation bout. At Kitanglad, only the female incubated the egg at night, but the male in Sinaka did so in three occasions. Females made short trips, averaging 0.32 hr (SE=0.18, n=3) to groom, defecate, collect sprigs, or interact with the male. The female did long trips as well that averaged 2.42 hr (SE=1.89, n=3) during periods when the male relieved her of incubation duties. For the females at the Tarragona and Kitanglad nests, long trips were made even when the male was not around. The longest time that the egg was left alone at Sinaka was 102 min, 73 min at Tarragona, and 111 min at Kitanglad (mean, 95.33; SD, 19.86). At Tarragona, the egg was exposed to 15 min of rain when the female was away for 73 min. The attending adults groomed, defecated, delivered sprigs, turned the egg, or fed in between incubation bouts. Grooming (i.e. preening, sunbathing, etc) was performed on either the nest tree or a nearby tree. Defecation was always in a different tree. Collected fresh branches were torn into pieces and deposited on the nest bowl. Before sitting, the egg was turned by arching the bill ahead of the egg and gently shoving it backwards towards the belly. Egg turning

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was seen at 29 instances between 0800-1600 hr at the Tarragona nests and was almost solely done by the female (93%). Incubating adults tugged at the nesting substrate and were seen dozing intermittently. During the incubation period, there were no attempts by the female to hunt. Table 5 Comparison of sex roles for the Incubation stage during daytime observations at three Philippine Eagle nests in Mindanao. Behavior
Nest attendance (%) Female Male Mean trip duration (hr) Female <1.0 hr >1.0 hr Male <1.0 hr >1.0 hr Inc. bout duration (hr) Female Male 1.43 (SD=0.81, n=70) 3.14 (SD=1.70, n=11) 1.00 (SD=0.87, n=33) 1.52 (SD=0.03, n=2)
th

Sinaka (N=18)
83 74 26

Tarragona (N=8)
82 95 05

Kitanglada (N=4)
78 100 0

0.15(SD=0.14, n=34) 4.59 (SD=1.72, n=8) 0.09 (SD=0.04, n=4) 21.04 (SD=3.82, n=5)

0.30 (SD=0.20, n=22) 1.08 (SD=0.10, n=3) 0.38 (n=1) >64.25 hr*

0.50 (SD=0.24, n=13) 1.60 (SD=0.35, n=2) b b 0.81 (SD=0.39, n=9) B

N = observation days, *male eagle attended nest only at the 8 observation day, a last four days of incubation, b = male did not attend to the egg during 4 observation days At the Tarragona and Sinaka nests, the male relieved the incubating female even without food. It appeared that a desire to incubate by either adult is signaled through vocalizations or by merely alighting near the nest. Mates seem to relinquish incubation by flying off the nest. Nest relief appears simple, with one bird walking into the nest bowl only when the other is already off. When inside

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the nest bowl, the attending adult approached the egg slowly with its toes clumped and claws bent inwards. During the whole observations when the female was incubating, the male hunted and delivered prey to the female. Prey delivery rate. Prey was delivered at a daily rate of 0.10, 0.25, and 0.71 at Sinaka, Taragona, and Kitanglad nests, respectively. It is remarkable to note that despite the low food delivery rate at the Sinaka nest (0.10), the pair successfully hatched an egg. At Tarragona, the egg was deserted. The Kitanglad nest had the highest prey delivery rate and breeding resulted to a successful fledging. It is suspected that the breeding failure at the Tarragona nest was a result the males failure to provide enough food to the attending female. The male did not bring prey for five consecutive days prior to egg desertion. The female made loud food begs but there were no response from the male. It is therefore not surprising that the female left and tried finding food for itself, to the peril of the egg. A similar incident happened at a 1983 nest whereby the female abandoned the egg and hunted when so little food was brought by the male (Lewis, 1983). In contrast, a high prey delivery rate during the incubation and throughout the nestling period (i.e., range = 0.710.61 prey per day) was sustained at the Kitanglad nest. Among several pairs of the Crowned Eagle in Kenya (Brown, 1966), a prey delivery rate of 1 in every 3.3 days (0.30 prey per day) was reported for the incubation period. This is lower than that recorded at the Kitanglad nest. However, Crowned Eagles hunt larger prey items so that lower prey delivery rates may be alright as long as hefty prey is delivered.

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3. Brooding Stage Sprig deliveries. For six consecutive days of observation of the Sinaka nest, a total of 18 sprigs were delivered, registering a daily rate of 3 sprigs. The female did most sprig deliveries (14 out of 18, or 78%). At the Kitanglad nest, the female solely delivered sprigs at a rate of 1.2 sprigs per day, with 81% of the sprigs (n=17 days) being fresh leafy branches collected from nearby trees. The average delivery rate at the two nests was 2.1 1.2 sprigs per day. Behavior on the nest. At the Kitanglad nest, the time spent for each bout of sitting on the chick was very variable (Table 5, range 7355 min). Shorter bouts and lower attendance rates were typical during fair weather, whereas longer bouts and higher attendance happened during heavy rains and cold, foggy weather. During one rainy day for example, the female sat on the chick for 355 min straight, emerging very drenched only when the rain stopped. Diurnal brooding ceased when the chick was 24 days old, but the female continued sitting on the chick at nighttime and stayed close during the day either on the nest or at nearby perches apparently to guard the chick. In both nests, diurnal and nocturnal brooding was solely by the female. The average attendance rate during brooding at two nests was 55.5%, which is lower than that for the incubation period (see Table 6). During brooding, the female also undertook two types of trips out of the nest; one lasting considerably less than an hour and another beyond an hour or more. Brief trips lasted 0.30 hr (SE=0.18, n=2) on the average, and were typically associated with self maintenance behaviors (e.g. grooming, sunbathing, defecation), sprig delivery

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and feeding. Female trips >1 hr averaged 1.47 hr (SE=0.26, n=2), and seemed to be associated with long breaks during a fine weather and probably hunting attempts too when the male had not delivered prey for some time. At Mt. Sinaka, for example, the female went out on a 3.82 hr excursion when the male has not delivered prey for 2 days. She came back with muddy talons and a soiled breast. At the Kitanglad nest, there were 17 instances where the female just stood near the chick, and the time spent doing this varied (range = 954 min, mean=0.35 hr, see Table 6). These bouts of standing coincided with sunny periods of the day and appear to help with shading the chick from the suns rays. In several instances, the female stretched its wings above the chick seemingly to shade the chick. Aside from shading and brooding the chick, the female also took the prey from the male and had the lead role in feeding the young. Table 6 Nest attendance, mean trip duration, brooding bout duration and standing bout duration during daytime observations at Sinaka and Kitanglad nests in Mindanao. Behavior Nest attendance (%) Mean trip duration (hr) <1.0 hr >1.0 hr Brooding bout (hr) Standing bout (hr) 0.17 (SD=0.19, n=12) 1.65 (SD=0.38, n=4) 1.34 (SD=0.76, n=14) 0.42 (SD=0.29, n=47) 1.28 (SD=0.16, n=6) 1.14 (SD=1.43, n=31) 0.35 (SD=0.28, n=17) 61 Sinaka (N=5) 50 Kitanglad (N=9)

N= observation days, SD=Standard Deviation

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Prey delivery rate. The prey delivery rate at the Sinaka nest was 0.50 prey per day, and the chick died because of exposure to very cold weather. In contrast, a delivery rate of 0.69 per day at the Kitanglad nest, which was sustained until the post-brooding period, resulted to a successful breeding. In terms of the frequency of delivery, the male at Sinaka delivered prey during the brooding period at a rate that is not very different from that at the Kitanglad nest (0.50 vs 0.69 prey per day). However, in terms of the average prey biomass delivered, the male at Sinaka did poorly. Within nine days after hatching, the three prey items brought by the male, which consisted of two rodents and a hind quarter of a flying lemur, only amounted to a rate of 269.17 g per day (Table 7). In contrast, the male at the Kitanglad nest delivered prey at a rate of 1,823.75 g per day. Hence, what the male at the Sinaka nest brought to the nest were very much insufficient to satiate both the female and the chick. The female at Sinaka left the nest for almost four hours in an apparent attempt to hunt, exposing the chick to rains and strong winds. Whole broods of Sparrow hawks Accipiter nisus in Scotland have suffered similar mortalities from sudden downpours when the female was away hunting (Newton, 1979). All of these underscore the importance of either frequent provisioning, if the prey items involved are small, or deliveries of hefty prey at reasonable intervals to prevent egg and chick abandonment during the critical stages of incubation and brooding stages.

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4. Post-brooding stage Sprig deliveries. The first 67 days of the nestling period registered the highest sprig delivery, at a rate of 1.1 sprigs per day ( 0.1, n=3) at the Kitanglad nest, and all were made by the female. Deliveries waned during the latter part of the post-brooding stage (Fig. 2). Almost all of the sprigs were fresh (85%, n=48), with dried sprigs (7) brought only at the latter part of the post-brooding stage. Sprig delivery stopped altogether when the eaglet was 107 days old (i.e. 3 months and 17 days). At the Cabuaya nest, sprig delivery was never seen during the 13 days of observation. At this time, however the estimated age of the young bird was four months and the adults were seen only on the nest during prey delivery and feeding.

Fig. 2. Sprig delivery rate at Kitanglad and Apo nests during the post-brooding period in Mindanao. 18 = 18-27 days, etc.

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At Apo, the first two months of the nestling stage was missed. Using photos of captive-bred chicks and by backtracking from when incubation and nestling was first verified, it was estimated that the Apo eaglet was about 2 months old at the start of the study. Sprig delivery rates at Apo seemed lower when compared with the same age bracket for the Kitanglad nest (Fig. 2). More than three-quarters of the sprigs delivered were fresh (19, n=24) and the female brought twice as many sprigs (67%, n=24) as the male. But unlike at Kitanglad, sprig delivery at Apo continued all through out the nestling stage. In both nests, however, sprig delivery was mostly made during early morning (i.e. 0500-0900 hr, Fig. 3).

Fig. 3. Times of the day sprig was delivered at Kitanglad and Apo nests during the post-brooding period in Mindanao. 5 = 0500-0559 hr, etc.

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It is interesting to note that the delivery of sprig to the nest continued from the nest-building phase until the post-brooding phase. Because the delivery of nest materials continued across the nesting season, investigators believe that nest-building has other functions. Besides the mechanical function of consolidating the nest, sprig deliveries are thought of as (1) a means of nest sanitation by covering debris on the nest, (2) a signal to advertise occupancy of a nesting territory, (3) and/or a means of maintaining humidity in the nest (as discussed in Newton 1979). Another explanation is that the use of green plant materials serves to repel ectoparasites harmful to the young (Wimberger, 1984). For Philippine Eagles, the nest sanitation hypothesis appears weak, because instead of covering uneaten prey remains, the parents remove them from the nest. This is obviously a quicker and more effective way of nest sanitation to avoid ant attacks, fly infestation, and the endoparasites they can bring. The occupancy hypothesis is equally weak, as Philippine Eagle breeding densities have been reported to be very low (70130 km2/pair, see Bueser et al, .2003) so that competition for territories might not be as intense as in other birds of prey. Maintaining humidity is unlikely a reason as humidity is not a problem in a tropical forest. The ectoparasite-repellent theory, in the absence of other probable reasons, might be true for Philippine Eagles, and is a good subject for further investigation. It would be interesting to know as well if eagles gather sprigs only from specific trees. One can also study the rate of decay of sprigs deposited to the nest bowl and see if these correlates to the frequency of sprig delivery.

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Adult behavior on the nest. Eagle behavior was observed almost throughout the post-brooding stage at the Kitanglad nest. At 39 days into the nestling period, the female delivered its first hunt after a day of absence. Although occasionally watched over from a distant perch, the chick was left alone on the nest even when there were rains beginning 39 days. Nighttime brooding stopped altogether as well. The chick was only attended to when fed. The female also spent substantially more time out of sight presumably hunting, which was the case also at the Apo and Cabuaya nests where the chicks were already old when found. The male still transferred prey to the female, but when the female was not around, the male fed the chick himself. Both the female and male occasionally ate from the prey while they feed the young. Prey delivery rates. Overall, a mean of 0.71 (SD = 0.1, n=3) prey item per day of observation was delivered during the post-brooding phase. On a per nest basis, the daily rate for the delivery of prey individuals was highest at the Apo nest (0.81 animals/day), followed by the Cabuaya nest (0.69 animals/day), and the Kitanglad nest (0.67 animals/day) (see Table 7). In Apo, there was one case where three animals were delivered in one day (two by the male), 9 instances where prey was rationed by each adult on the same day (the female doing so first on 6 occasions), and two instances where the male alone carried prey twice in a single day. The shortest and longest delivery interval between male and female was 48 min and 321 min, respectively (mean = 181.9 110.1, n = 9 obs). The mean gap for exclusively male deliveries is 349.5 min 30.4.

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Multiple deliveries also occurred at the Kitanglad nest by the pair and the male by itself, each on three occasions. The shortest and longest interval in male and female delivery was 35 min and 445 min, respectively (mean = 286.7 220.4, n = 3 obs). The shortest gap for exclusively male delivery was 60 min and the longest was 244 min (mean = 135 95.1, n = 3 obs). In contrast, no multiple deliveries were observed at the Cabuaya nest. Table 7 Prey delivery rates during the brooding and post-brooding (early and late) stages of the nesting period in three Philippine Eagle nests in Mindanao.
Nests
Brooding ( 28 days) No./ day Sinaka combined female male Cabuaya combined female male Apo combined female male unknown Kitanglad combined female male 0.69 0.69 1823.75 1823.75 0.86 0.48 0.38 1205.24 780.48 424.76 0.56 0.12 0.44 1131.76 287.79 843.97 0.67 0.25 0.42 1159.82 475.91 683.91 0.89 0.44 0.33 0.11 1002.30 399.52 534.63 68.15 0.78 0.41 0.37 892.25 399.68 492.57 0.81 0.42 0.36 0.03 924.86 399.63 505.19 20.04 0.69 0.38 0.31 1803.85 825.38 978.46 0.69 0.38 0.31 1803.85 825.38 978.46 0.50 0.50 269.17 269.17 Biomass/ day (g) Early postbrooding (29-80 days) No./ day Biomass / day (g) Late postbrooding ( 81days) No./ day Biomass/ day (g) No./ day Biomass/ day (g) Post brooding

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However, if we look at the average biomass of prey delivered each day, Cabuaya had the highest rate (1803.85 g per day) and Apo had the lowest (924.86 g per day). As shown in Table 7, adults at the Cabuaya and Kitanglad nests hunted heavier prey items across the post-brooding stage while adults at the Apo nest took lighter prey items. At the Apo and Cabuaya nests, the female delivered more prey individuals than the male, but in the Kitanglad nest, the male substantially brought more than the female (Fig. 4). Looking at the average prey biomass though, the male consistently brought heavier prey animals in all of the three nests than the female (Fig. 5).

Fig. 4. Comparison of the mean (n=3) for male, female and combined malefemale values for the number of prey individuals delivered per day to similar measures for each of three nests (Cabuaya, Apo, Kitanglad) in Mindanao.

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Over-all, prey delivery rates at the Cabuaya, Apo and Kitanglad nests during the post-brooding stage were relatively high (Table 7). At this stage, the female had stopped brooding and guarding the young and was already helping out with hunting. This is consistent with reports that food deliveries become more frequent at the latter stages of the nestling period where the food demands of a growing chick are greater (Newton, 1979). All attempts at the three nests resulted in a successful fledging. Hence, an average prey delivery rate of 0.72 prey per day and 1,823 g. per day in terms of biomass during the whole nestling period can be the average, if not the minimum, to ensure a successful nesting attempt.

Fig. 5. Comparison of the mean (n=3) of male, female and average biomass delivered per day, with similar values for each of the male, female, and combined rates at each of the three Philippine Eagle nests observed (Cabuaya, Apo, Kitanglad) in Mindanao.

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Diel pattern of prey delivery. At the Cabuaya nest, prey delivery during the post-brooding stage followed a strongly bimodal pattern throughout the day. The largest peak in deliveries occurred at mid-morning (0800 to 1000 hr), with a second peak in the afternoon (1400 to 1600 hr). A similar pattern was also observed at the Kitanglad nest. At the Apo nest, however, prey delivery rose steadily throughout the whole morning until early afternoon, decreasing in late afternoon. Pooled data for the three nests retains a bimodal pattern in the timing of prey deliveries to the nests (Fig. 6).

Fig. 6. Diel pattern of prey delivery at 3 Philippine Eagle nests (Cabuaya, Kitanglad, Apo) during the post-brooding period in Mindanao. 6 = 06000759 hrs, etc.

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Looking at the diel pattern of delivery according to prey weight (Fig. 7), the bimodal peaks in delivery remained evident. The peak of delivery of prey within the 1-499 g range seems to occur between 0800-1000 hr, and waning thereafter. Delivery of long-tailed macaques (i.e 4000 g) had two peaks, one between 0800 to 1000 hr and between 1200 to 1400 hr. For flying lemurs (i.e. 1000 g), more were delivered during early morning and around noon. No prey items from the 2000 g category were brought to the nest. Over-all, prey items within the 500 g category had the highest frequency.

Fig. 7. Diel pattern of delivery according to weight of prey at 3 Philippine Eagle nests during the post-brooding period in Mindanao. 1 = 1 499 g, etc.

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5. Parental Care Philippine Eagles have the longest and energetically most expensive parental investments for any birds of prey (see Newton 1979). They consistently rear only a single young within two full years for an entire breeding cycle (Collar et al., 1999). Courtship and nest-building until egg-laying lasted four months at the Sinaka nest. Incubation lasted 58 days at the Sinaka nest and 6061 days at the Davao del Sur nest (Gonzales 1968). In captivity, eggs hatch after 5657 days of artificial incubation (Tadena et al., 1999). Among large raptors, only the Andean Condor incubates eggs for at most 58 days (Brown and Amadon 1968). At the Kitanglad nest, the nestling period lasted 146 days, which is within the range of 104164 days as observed in previous studies (Gonzales 1968, Kennedy 1985). Parents continued feeding the young until the next nesting season. In case of breeding failures during the first quarter of the year, they renest towards the end of the year. This occurred at the Kitanglad nest wherein the chick studied was a product of a re-nest in December 2006 when an 89 month old eaglet of the same pair was shot and removed from the nest site in September 2006. At the Tarragona nest where breeding failed in January 2007 a re-nest happened December of the same year. Several re-nesting after a failed attempt were also documented in at least four nesting territories monitored in Mindanao since 1977 (Miranda et al., 2000). Male and female roles during the nesting season were consistent with previous reports (Gonzales, 1968; Kennedy, 1985). The primary role of the male is to acquire food during the incubation and nestling period. The male also

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relieved the female from incubation duty and fed the chick as well, but only rarely. The primary role of the female is to incubate the egg and brood, guard and feed the nestling. Females never hunted during incubation, or if they did, during long forays away from the nest, there were no recorded successes. Parental care in Philippine Eagles, whereby males provide nourishment to the female and the young, and commonly play very little role in incubation and feeding the young, conforms to that seen in the Harpy Eagle Harpia haypyja, its ecological equivalent in South America (Rettig 1978), and to other raptor species as well (Newton 1979).

E. Nestling Development and Behavior

Almost the whole nestling period was documented at the Kitanglad nest. Hatching was not actually seen, but was inferred when the female seemed to sit lightly on the nest bowl on January 21. Feeding was first seen on the same day, but it was not until two days later that the chick was seen. By day 4, the chick began crawling about. By day 12, the chick could sit, stand and swallow larger pieces of meat, including the metatarsals of a palm civet. By 24 days, the chick had gained its daytime thermal independence and could already walk well around the nest bowl, doing quick wing exercises and playing with sticks, leaves and prey remains (i.e. object play). By 43 days, it could feed by itself although the adults fed it throughout the nestling period. At 54 days its behavioral repertoire had widened to include flight exercises, wing-slaps (i.e. single loud wing flaps) and mock attacks at prey remains and sticks on the nest which intensified as the

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chick grew older. However, most of the day was spent on general maintenance behavior, but mostly either just standing or prostrate (brooding or sleeping) (Fig. 8). Generally, older chicks at Cabuaya and Apo nests behaved similarly (Fig. 8).

Fig. 8. Percentage of time spent on diurnal activities by the nestling during the post-brooding behavior at the Cabuaya, Apo and Kitanglad nests in Mindanao.

Within 3 weeks before fledging at the Kitanglad nest, the nestling explored adjacent branches. At this time, agility exercises such as running across the nest and wing-flapping while jumping were frequent. The chick also did hop-flights to and from nest-support branches. Similar excursions and hop-flights and

increased frequencies of flapping exercises were seen in the chicks at Apo and Cabuaya. The nestling also grabbed prey from attending adults and called vigorously at adults on sight. In one occasion at the Kitanglad nest, the chick was

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very aggressive towards the male on the nest while it was feeding on the prey delivered by the female. The male was forced out of the nest by the vigorous calling and aggressive advances of the young. The chick flew out of the nest when it was 146 days old.

F. Diet and Food Habits 1. Prey Species and food habits

Taking all five Philippine Eagle pairs together, we recorded a total of 151 prey items (107 identified to the species level) from pre egg-laying until the postbrooding periods (Table 8). At least 14 taxa were recorded and 9 were identified to the species level. Nine of the prey taxa were mammals, which comprised 91% of identified prey items. The top two prey items in terms of percentage frequency were the Philippine flying lemur Cynocephalus volans (29.8) and the Common palm civet Paradoxurus hermaphrodites (13.9%). Both species are arboreal and nocturnal. Another nocturnal and arboreal prey item is the Mindanao flying squirrel Petinomys crinitus, and it ranked fourth in terms of percentage frequency (5.3%). The Long-tailed macaque Macaca fascicularis, an arboreal mammal that is active only during the day, was the third most important prey item (10.6%). However, if we look at which prey species had the highest biomass contribution, the long-tailed macaque (34.8%) substantially becomes the most important prey.

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Table 8 Quantity of Philippine Eagle prey species observed directly from courtship through the nestling stages at four nest sites in Central and Eastern Mindanao, Philippines, from 1999 to 2007. Species Mammals Cynocephalus volans Paradoxurus hermaphroditus Petinomys crinitus Macaca fascicularis Cannis familiaris domesticus Felis domesticus Unidentified mammal Unidentified rodent Unidentified bat Reptiles Naja samarensis Tropidolaemus wagleri Unidentified snakes Birds Sarcops calvus Gallus gallus domesticus Total Total mammals Total birds Total reptiles No. Frequency (%) 30.0 14.0 5.3 11.3 2.0 1.3 14.7 8.7 4.0 Biomass (g) 61200 67200 1800 85000 1500 1000 5500 3315 186 % Biomass 26.2 28.8 0.8 36.5 0.6 0.4 2.4 1.4 0.1

45 21 8 17 3 2 22 13 6

1 3 3

0.7 2.0 2.0

500 600 600

0.2 0.3 0.3

1 6 151 137 7 7

0.7 4.0 100.0 90.7 4.6 4.6

170 4602 233173 226701 4772 1700

0.1 2.0 100.0 97.2 2.0 0.7

Almost all of the prey animals brought to the nest were decapitated and dismembered. Typical for almost all of the mammalian prey, only the torso and the hind-quarters were intact. All were apparently from fresh kills. In the Kitanglad nest for example, a Philippine flying lemur was delivered to the nest alive, and in the Apo nest, a gravid pit viper was still twisting and struggling against the adults

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talons when it was fed to the chick. Almost all parts of the prey items brought to the nest were consumed, including entrails and whole limbs. Fore and hind limbs were swallowed whole by the adults and the eaglet as well when it was older. Uneaten prey remains such as dried flying lemur skin and a leg of a palm civet were removed from the nest. Our observations on the prey delivered to nests revealed both similarities and differences to those reported by previous investigators. Mammals dominated the diet as has been found by previous studies (Gonzales, 1968; Kennedy, 1985) and flying lemurs were similarly the most important prey species by frequency. But in terms of biomass contribution, this study showed that long-tailed macaques became the most important prey species. In contrast, macaques are a minor prey at the Davao del Sur nest in 1963. Out of the 48 prey items recorded, only three were monkeys (20% of the total biomass). Flying lemurs comprised 79% of the total biomass (Gonzales, 1968). At three nests studied by Kennedy (1985), there were 19 times more flying lemurs than macaques, making the only primate in the country again a very minor prey. The fact that more of a particular prey is delivered to one nest but fewer of it at a different nest only reinforced the previous hypothesis that Philippine Eagles are opportunistic feeders, taking whatever is available and easy to catch (Kennedy, 1985). Such ability to shift diet depending on prey availability was also observed among Crowned Eagles and Harpy Eagles. Crowned Eagles prefer primates in forest habitats, but hyraxes and duikers in

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open savannas (Mitani et al., 2001). Harpy Eagles favor primates in one territory, but sloths in another (Galetti and Carvalho, 2000). The high proportion of nocturnal mammals indicates the importance of arboreal and nocturnal animals in the Philippine Eagle diet. Flying lemurs are gliding foliagivores. They forage by night and roost during the day either in tree cavities or by gripping a trunk or a branch with its patagium cloaking its body (Wischusen and Richmond, 1998). The common palm civet is also arboreal and rests up in the trees during the day in hollows or shelter of entangled vines (Shiroff, 2002). There is no information on the roosting habits of flying squirrels, but they could be cavity roosters as well, like the rest of the flying squirrels (Fox and Mulheisen, 1999). These species are found from sea level up to the mountains, but they are particularly common in the mid-elevation to montane forests (Heaney et al., 1998). Because they seem to be thriving well in a habitat that is not under immediate threat, arboreal mammals will most likely remain as an important component of Philippine Eagle prey base across Mindanao. A diet study of flying lemurs by Wischusen (1990) showed that they do not feed on leaves of dipterocarp trees. The trees they feed on are common in midelevation to montane forests. Flying lemurs is not a threatened species as well. Because the flying lemur population in Mindanao appears stable (Heaney et al.,2006) and its diurnal roosting habits seem to make it more vulnerable to eagle predation, the flying lemur will most likely remain as the prime prey item in Mindanao.

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In studies of large tropical raptors like the Harpy Eagle and African Crowned Eagle, prey remains collected on the nest and at the base of the nest tree is a major source of diet information (Rettig, 1978; Galetti and Carvalho, 2000; Mitani et al.,2001). In contrast, in the case of the Philippine Eagle where prey is dismembered, bones swallowed and the skull of large prey items discarded elsewhere, collection of prey remains on the nest and under nest trees to study diet is not appropriate. Direct observation of nest and probably video recordings from hidden nest cameras that are installed way before the egg laying period appear to be the best means to study and quantify diet.

2. Philippine Eagle hunting techniques

The wide array of prey items that resulted from studies by Gonzales (1968), Kennedy (1985) and this study highlights the diversity of hunting styles employed by the Philippine Eagle to catch prey. Predation on nocturnal mammals seems to entail aptitude at tracking roost sites or tree cavities from which to pick off resting animals. The relatively slender legs of the Philippine Eagles have been speculated to be an adaption to taking out mammals inside tree cavities (Kennedy, 1981). For macaques, deer, hornbill and snakes, slyness, speed and an element of surprise appear crucial. Crowned Hawk-eagles (Mitani et al.,2001) sit and wait in the canopy or swoop down from high to take on monkeys. Philippine Eagles appear to employ the same method. Kennedy (1977) documented direct pursuit of prey as a hunting strategy, which seems important for hunting along valleys where visual track of prey animals can be maintained.

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3. Predation on domestic animals Domestic animals as prey items were recorded at the Apo nest. The domestic chicken Gallus gallus bankiva was relatively the most represented (n=6), and was almost exclusively brought in by the female (n=5). The male was solely responsible for the two records of domestic cats Felis domesticus on the nest, but contributed only one of three puppies Canis familiaris brought to the nest. Together, domestic prey represented 16% of the total identifiable prey at the Apo nest. In the pooled data, however, domestic animals represented only a mere 7.3% of prey items, and contributed only 3.0% of the total prey biomass. These observations represent the first confirmed record of domestic prey brought to the nest by Philippine Eagles. The first published report of domestic animal predation by Philippine Eagles was perhaps by Ogilvie-Grant in 1897 when he wrote that it visits the villages and carries off domestic poultry and Whitehead in 1899 when he said that the Philippine eagle is well known to the natives as a robber of their poultry and small pigs (in Collar et al., 1999). Gonzales (1968) commented on domestic predation at a nest in Davao del Sur: a native womanasked me to reimburse her piglet, allegedly taken by one of the parent eagles. The study of the Apo nest provided the first evidence for domestic animal predation in Philippine eagles, with cats, puppies and domestic chicken altogether representing 16% of the total prey items. No domestic animals were ever brought to the four other nests, nor were there one in the single nest studied by Gonzales (1968) or the five nests studied by Kennedy (1985). Upland residents consistently report heavy

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predation on domestic animals, and often justify shooting or trapping of Philippine Eagles as a way of alleviating their losses. Although we got evidence from a single nest, it is important to look further into the occurrence of domestic predation especially since Philippine Eagles (as will be discussed further later) has the ability to shift its preference of prey to whatever is available and easier to catch and handle. Philippine Eagles do take domestic prey, and an interesting challenge that comes out from this result is how to convince the public not to persecute the birds for taking their livestock and pets. Denying such knowledge to the public is out of the question, as this would certainly cause more harm than benefits. From the viewpoint of a resident who has personally experienced livestock or pet loss from the eagles for example, continued denial can lead to feeling of animosities not only to the bird but also to conservation practitioners. It can potentially breed more hate than care and sympathy for the bird and the cause to conserve it. In the uplands where opinions are easily taken for facts, providing the right information and transparency is critical. An effective strategy to address people and eagle conflict as a result of domestic predation can be drawn from the nest site at Apo. Reports of domestic predation were common long before this study. In 1996, a community-based conservation (CBC) program was launched by PEF which included setting up and strengthening a local cooperative, training and material assistance for sustainable upland agriculture and marketing, soft financial loans, and provision of a marketing vehicle. That project was fully turned over to the community in

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2002, and the pair has continually bred since then in this disturbed nesting site, with another young successfully fledged in March 2008. Despite the occasional domestic predation which probably happens with each nesting cycle, five successive nesting attempts were recorded since 2003, and resulted in the successful fledging of 3 young (60% breeding success). As a result of investing in CBC at Apo, the community maintains a positive attitude towards the eagles and has considered the occasional loss of livestock as their generous contribution to an imperiled eagle neighbor. CBC approaches are a tested tool for effective species and habitat conservation (Horwick and Lyon, 2007), and should be implemented in each nesting territory where human-eagle conflicts exist.

4. Philippine Eagle diet and food habits at Apo and Kitanglad nests

Substantial prey information was recorded from the Kitanglad and Apo nests where a lot of observation efforts were spent. Interestingly, the nest differed in terms of prey diversity. Prey items at the Kitanglad nest were all mammals, whereas reptiles and birds formed part of the diet at Apo in addition to mammals. The pair at Kitanglad had a narrower Food Niche Breadth (FNB, 3.93) than at Apo (8.31). However, the estimated Mean Weight of Prey (MWP) delivered was heavier at the Kitanglad nest than at the Apo nest. The average prey biomass per day delivered at the Kitanglad nest is heavier and consisted mostly of medium to large mammals that accounted for a heftier biomass (Table 9). On the other hand, the prey brought by adults to the Apo nest included lighter reptiles and birds. Dietary overlap between the two pairs was 0.61.

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Table 9 Food-niche breadth (FNB), dietary overlap, and estimated mean weights (g) of prey of Philippine Eagle pairs at Kitanglad and Apo during the nesting season in Mindanao. Food-niche parameters Total prey individuals Mammal total individuals Bird total individuals Reptile total individuals Mean weight of prey (g) FNB Dietary overlap 1758.94 3.93 0.61 Kitanglad 52 52 Apo 73 60 7 6 1107.23 8.31

The difference in the prey composition in the diet between the Kitanglad and Apo eagle pairs illustrates the breadth of prey that the eagle can take. An overlap index of 0.6 indicates diet similarity, but also a certain level of difference that is worth looking at. Six taxa were unique to the Apo nests, namely bats, rodents, snakes, birds, domestic cat and domestic dog. Based on mean prey weight calculations, the Kitanglad pair went for heavier prey items. Contrasting the habitat types at these two sites, one can not help but ask whether such differences in prey taken were a product of habitat quality in these two sites. For example, the Apo nest site is very much disturbed whereas the Kitanglad gets very little human interference. Kitanglad has a greater mammalian diversity (Heaney et al., 2006) and has better protection and management measures than Apo. The fact that the pair at Kitanglad specialized on a few native and heavier mammalian prey while the pair at Apo used a range of vertebrate groups,

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including domestic animals, points to the possible relationship between habitat quality and prey preference in Philippine Eagle pairs. It is hypothesized that eagle pairs in large and undisturbed habitats tend to specialize on moderate to large arboreal mammals, whereas those in disturbed areas have a broader spectrum of vertebrate prey, including animals associated with highly disturbed areas. Corollary to this hypothesis, it also predicted that whenever heavier arboreal mammals are in low densities, the pair would compensate by taking on more of a wide variety of smaller prey items.

5. Size difference between prey items hunted by male and female eagles

The size of prey delivered by female and male eagles varied across the post-brooding phase (Fig. 9), although pooled data indicated that the females delivered more of the smaller prey (i.e. weighing less than 1 kg). In contrast, males delivered more animals within the 1,000 to 4,000 g range. During the brooding phase at the Kitanglad nest, the male exclusively brought prey items within the 1,000 to 4,000 g range. These animals were the Philippine flying lemur (n=2), Common palm civet (n=2), and Long-tailed macaque (n=2). Calculating the total biomass of prey at the three nests, the male had a bigger share (91,888 g). The sum of the females hunts (66,921 g) is only 72% of the total biomass contribution of the males.

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Fig. 9. Comparison between the prey mass categories brought by male and female during the post-breeding phase in three Philippine Eagle nests observed (Cabuaya, Apo, Kitanglad) in Mindanao. 1= 1499 g, etc It has been hypothesized that the size difference among raptors where the females are lager than the males (i.e. reversed sexual dimorphism) is an adaption so each sex can exploit different prey species (Selander, 1969). Another hypothesis suggests that smaller males make hunting of small and more abundant prey more efficient, whereas larger females can take on larger prey to help meet the food demands of a growing chick during the latter stage of nesting (Reynolds, 1979). Looking at the biomass of prey brought to three nests, both males and females hunted large and moderate-sized prey items. In fact, the male brought in more of the heavier prey items on the average (e.g. macaques, civets

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and flying lemurs) than the female, and the female seemed to have specialized on smaller prey items than the male. At two nests, prey brought during the postbrooding period was likewise variable, with males also bringing in heavier prey than females. Thus, these results do not seem to support the prey augmentation hypothesis mentioned. Sexual dimorphism among Philippine Eagles, whose diet is mainly mammals, can be considered moderate. Calculating for its dimorphism index by getting the ratio of female to male wing length, the value for Philippine Eagles is about 1.3 (Appendix H). Raptors of the genus Falco and Accipiter exhibit moderate to extreme dimorphism with an index ranging from 1.101.70 (Appendix I, data from Clark, 1999). According to Newton (1979), raptors that feed on fast and agile prey, like birds, exhibit stronger dimorphism than raptors that either exclusively feed on mammals or raptors that have a bird-mammal combination in their diet. The extent of dimorphism among Philippine Eagles seems to conform to this general trend in diet and dimorphism.

6. Breeding failures as an effect of food stress

This study showed two possible ways by which low food delivery rates can lead to breeding failures: one is egg desertion and the other is nestling desertion. In both cases, the egg/chick becomes vulnerable to predators and the elements in the absence of the attending female. The study also demonstrated that prey biomass rather than the number of prey individuals becomes more important in assessing how much nourishment is necessary to ensure successful

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reproduction. The abundance and availability of prey is critical to the breeding and survival of the Philippine Eagle. In fact, several investigators showed that the patterns of habitat use by raptors are very much dependent on the distribution of prey within such habitat (Newton, 1979; Ward et al.,1998). However, the influence of prey density and availability on Philippine Eagle ecology, behavior and productivity has not been given the rightful research attention it deserves as far as studying prey base is concerned.

G. Estimates for Philippine Eagle adult longevity Using Browns (1966) formula, it is estimated that in a stable population an adult Philippine Eagle must live 29-30 years on the average before it can breed a young that will reach sexual maturity (Appendix G). This age is not impossible because a captive female at the Philippine Eagle Center is 46 years old now and still alive. However, with the occurrence of human-induced mortalities which might already be causing recent instability in the eagle population, the calculated longevity estimate might represent a minimum value only. Preventing high death rates among breeding adults, therefore, is very critical in the light of the estimated longevity of adult Philippine Eagles. If many adults die before reaching the average adult longevity, few young eagles will be produced to replace dying adults. Coupled with high juvenile mortality, continuing damage to nest sites, declining prey base, deforestation, and widespread apathy of Filipinos to the plight of the species, the eagles will inevitably fall into the vortex of extinction.

Chapter 5

SUMMARY, CONCLUSION AND RECOMMENDATIONS

Summary This thesis described the initial results of a continuing effort to document the nest and nest tree characteristics and the breeding success, biology, behavior and food habits of nesting Philippine Eagle pairs studied in Mindanao from 1999 to 2007. Baseline values for breeding behavior parameters, (i.e., nest attendance, incubation and brooding bout duration, trip duration and feeding rates during each stage of breeding) were provided. The preliminary estimate of longevity illustrated the critical value of ensuring the long-term survival of breeding adults to maintain population stability. Results for breeding behavior and food habits were not very different from previous studies, but this study provided details and additional insights. Flying lemur (Cynocaphalus volans) is the primary prey species, and arboreal mammals as the most important prey group. Because arboreal mammals appear to exist in stable numbers in mid-elevation forests that are not immediately under human threat, these animals will most likely remain as an important prey group in Mindanao. Philippine Eagles can shift their diet to what is available. When heavier, native mammals are scarce, they seem to adjust by taking on more of smaller prey items. In contrast to the prediction that females in sexually

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dimorphic raptors tend to take on large and heftier prey than males, the reverse seems to be true for Philippine Eagles. This study also provided the first clear evidence of predation on domestic animals and pets by Philippine Eagles. Because they are opportunistic feeders and because domestic predation may result to eagle mortalities if people shoot the birds to alleviate their losses, predation by Philippine Eagles needs to be investigated further. The study also provided insights into the status of nest sites and breeding pairs and gaps in research information that needs to be addressed. Because it is long-lived, has an average longevity of 29 to 30 years, and is very dependent on a tropical forest that is fast-disappearing, the Philippine Eagle is very vulnerable to extinction. With the loss of large trees in lowland forests,

many nest sites are concentrated along forest edges near upland communities, making nesting adults vulnerable to human persecution. Being opportunistic feeders, they are predisposed to capturing domestic prey when native prey is limiting. Predation on domestic animals makes the eagles more susceptible to shooting and trapping. There is evidence that some pairs suffer from lack of food that may result to breeding failures through egg or nestling desertion. But there is evidence too that eagle pairs near large, relatively intact forests where stable populations of arboreal mammals exist are breeding well. Prey abundance and availability is important for the long-term survival of the Philippine Eagle. Because of the importance of increasing adult survival as well as ensuring productivity, protection of breeding Philippine Eagles and the places where they

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nest are important. Conservation of the home range where they forage is equally important to ensure that enough prey base is available. Community-based conservation (CBC) approaches can be a potent tool for conservation in places where people and Philippine Eagle conflicts are tense.

Conclusion This study provided evidence that the Philippine Eagle has the longest and energetically most expensive parental care for any birds of prey. Because of its breeding biology and the fact that the species is long-lived, it is very vulnerable to extinction. This study supports a previous suggestion that breeding failures are not the primary cause of decline. In the light of the trends in human persecution, deforestation and the fact that the species is long-lived, adult mortalities seem to be the major cause of decline. Philippine Eagles in Mindanao nest exclusively on large trees, which could already be limiting. Some Philippine Eagles have no choice but to nest at forests edges near communities where most large trees are confined. Patterns of courtship, aerial displays, nest-building, and copulation rates are similar to those known for other birds-of-prey. Philippine Eagles invest on frequent copulation perhaps to strengthen pair bond, and on sprig delivery all through out the nesting season probably as a means to rid the nest of ants, flies and ectoparasites.

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The flying lemur (Cynocephalus volans) remains to be the dominant prey item, and arboreal forest mammals as the most important prey group in Mindanao. Philippine Eagles are opportunistic feeders. Initial data suggests that pairs in disturbed habitats feed on more prey species that weighed less whereas those in intact, less disturbed forests feed on fewer but relatively heavier prey. Philippine Eagles feed on domestic animals, but this maybe happening only in heavily disturbed nesting territories where natural prey items are scarce. Predation of Philippine Eagles on domestic animals and pets has conservation implications as it makes the birds even more susceptible to hunting or shooting. Each breeding adult must live for an average of 29 to 30 years so it can produce a young that would replace it. Conservation strategies that increase adult survival are important to population stability.

Recommendations 1. Information from birds nesting in forest interiors must be collected. The information provided in this study is limited to observations of pairs in forest edges, and may not accurately represent the Philippine Eagle population as a whole. There could be eagles nesting at the north and north-east end of Mindanao, particularly at Mt Hilong-hilong, Pantaron Range, and the forests of Lanao Provinces, where huge tracts of lowland forest are still found. 2. More ecological information is needed from pairs on other islands (i.e. Luzon, Leyte, and Samar islands) to determine general patterns and as well as uniqueness in island populations.

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3.

Comparative studies that look at behavior, productivity and survival between edge-nesting and interior-nesting eagles, as well as between birds in small patches against those in larger forests must be conducted.

4.

More prey base studies are needed, including assessing the relationship of prey abundance and availability to home range size, movement patterns and breeding success.

5.

Radio and remote telemetry techniques must be employed to measure habitat use, movement and dispersal of adults and dispersing young.

6.

To complete our understanding of Philippine Eagle breeding biology and ecology, studies of the post-fledging stage must also be done.

7.

A study of nest site selection and GIS mapping and modeling of actual and potential nest site must be done to know if nest sites are indeed limiting.

8.

Adult and young eagles must be marked and banded as a means to know mortality or turn-over rates.

9.

Long-term monitoring of breeding in nest sites must be sustained.

10. For current conservation efforts, species-directed approaches much invest on minimizing, if not preventing, adult mortalities. 11. For Philippine Eagles nesting in forest edges close to communities, Community-based Conservation approaches must be implemented. 12. On-site education campaigns must be designed to prevent destruction of nesting sites and shooting and trapping of adult eagles and their fledged young.

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Davies, R.A.G. (1999). The extent, cost and control of livestock predation by eagles with a case study on Black Eagles (Aquila verreauxii) in the Karoo. J. Raptor Res, 33, 67-72. del Hoyo, J., Elliott, A., & Sargatal, J. (Eds). (1994). Handbook of the birds of the world. Vol. 2. New world vultures to guineafowl. Barcelona: Lynx Edicions. Dunning, J.B. Jr. (Ed). 2000. CRC Handbook of Avian body masses. Florida, USA: CRC Press. Fox, D., & Mulheisen, M. (1999). Animal Diversity Web Glaucomys volans. http://en.wikipedia.org/wiki/Southern_Flying_Squirrel. [Accessed March 30, 2008]. Galleti, M., & de Carvaljo, O. (2000). Sloths in the diet of a Harpy Eagle nestling in Eastern Amazon. Wilson Bull., 112(4), 535-536. Gonzales, R.B. (1968). A study of the breeding biology and ecology of the monkey-eating eagle. Silliman Journal. 15, 461-491. Gonzales, R.B. (1971). Report on the 1969 status of the monkey-eating eagle on Mindanao Island, Philippines. Bull. Int. Coun. Bird Preservation, 11, 154168. Grossman, M.L., & Hamlet, J. (1969). Birds of prey of the world. New York: Clarkson N. Potter, Inc. Harvey, P.H., Greenwoodan, P. J., & Perrins, D.C.M. (1979). Breeding area fidelity of Great Tits (Parus major). J. Anim. Ecol, 48, 305-313. Heaney, L.R., Balete, D.S., Dolar, M.L., Alcala, A.C., Dans, A.T.L., Gonzales, P.C., Ingle, N.R., Lepiten, M.V., Oliver, W.L.R., Ong, P.S., Rickart, E.A., Tabaranza, B.R.Jr., & Utzuurum, R.C.B. (1998). A synopsis of the mammalian fauna of the Philippine Islands. Fieldiana: Zoology, N.s. 88. Heaney, L.R., & Regalado, J.C.JR. (1998). Vanishing Treasures of the Philippine Rainforest. Chicago, IL: The Field Museum.

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Heaney, L.R., Tabaranza, B.R.Jr., Rickart, E.A., Balete, D.S. & Ingle, N.R. (2006). The mammals of Mt. Kitanglad Nature Park, Mindanao Island, Philippines. Fieldiana: Zoology, N.s.112. Holt, D.W. (1999). Frequency and timing of copulations in the Prairie Falcon. Wilson Bull., 104(2), 333-338. Horwich, R.H. & Lyon, J. (2007). Community conservation: practitioners answer to critics. Oryx, 41(3), 376-385. Jenkins, J.M., & Jackman, R.E. (1993). Mate and nest-site fidelity in a resident population of Bald Eagles. The Condor, 95, 1053-1056. Joseph, V. (1998). Emergency Care of Raptors. In Rupley, A.E.(Ed), The Veterinary Clinics of North America Exotic Animal Practice and Critical Care. Philadelphia, Pennsylvania: W.B. Saunders Co. Joshi, A., Smith, J., & Cuthbert, F. (1995). Influences of Food Distribution and Predation Pressures on Spacing Behavior in Palm Civets. Journal of Mammalogy, 76(4), 1205-1212. Kennedy, R.S. (1977). Notes on the biology and population status of the monkeyeating eagle of the Philippines. Wilson Bull. 89:1-20. Kennedy, R.S. (1981). Saving the Philippine Eagle. Natl. Geogr. 159, 847-856. Kennedy, R.S. 1985. Conservation research of the Philippine Eagle. Natl Geogr. Soc. Res. Rep, 18, 401-414. Kennedy, R.S., Gonzales, P.C., Dickinson, E.C., Miranda, H.C.JR., & Fisher, T.H. (2000). A guide to the Birds of the Philippines. USA: Oxford University Press. Kenward, R.E. (1999). Raptor predation problems and solutions. J. Raptor Res. 33, 73-75. Lack, D. (1940). Courtship feeding in birds. The Auk. 57, 169-178. Lehner, P. N. (1979). Handbook of ethological methods. London: Garland STPM Press. Lerner, H.R.L., & Mindell, D. P. (2005). Phylogeny of eagles, Old World vultures and other Accipitridae based on nuclear and mitochondrial DNA. Molecular Phylogenetics and Evolution. 37, 327-346. Lewis, R. (1986). A rain forest raptor in danger. Oryx, 20, 170-175.

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Lovell, H.B. (1947). Black vultures kill young pigs in Kentucky. Auk, 64, 131-132. Marti, C.D. (1987). Raptor food habit studies. In Giron-Pendleton, B.A., Millsap, B.A., Cline, K.W., & Bird, D.M. [Eds]. Raptor Management Technique Manual. Washington: Wildlife Federation. Matchett, M. R., & O'gara, B.W. (1987). Methods of controlling golden eagle depredations on domestic sheep in southwestern Montana. J. Raptor Res., 21, 85-94. Mcemeaney, T.P. & Jenkins, M.A. (1983). Bald eagle predation on domestic sheep.1983. Wilson Bull., 95(d), 694-695. Mcgahan, J. (1968). Ecology of the Golden eagle. The Auk, 85, 1-12. Miranda, H.C.Jr., Salvador, D.I., Ibanez, J.C., & Ibanez, G.B. (2000). Summary of Philippine Eagle reproductive success: 1978-1998. J. Raptor Res, 34(1), 37-41. Miranda, H.C.Jr., & J.C. Ibanez. (2006). A modified Bal-Cha Tri to capture Great Philippine Eagles for radio-telemetry. J. Raptor Res, 40, 235-237. Mitani, J.C., Sanders, W.J., Lwanga, J.S., & Windfelder, T.L. (2001). Predatory behavior of crowned hawk-eagles (Stephanoaetus coronatusI) in Kibale National Park, Uganda. Behav. Ecol. Sociobiol, 49, 187-195. Mock, D.W. (1985). An introduction to the neglected mating system. In Gowaty, P.A. & Mock, D.W. (Ed.) Avian Monogamy. Ornithological Monographs No. 37. Washington, D.C.:American Ornithologist Union. Newton, I. (1979). Population ecology of raptors. London: T. & A.D. Poyser. Pandolfi, M., Pagliarani, R., & Olivetti, G.. (1998). Intra-and extra-pair copulations and female refusal of mating in Montagus harriers. J. Raptor Res. 32 (4), 269-277. Phillips, R. L., & Blom, F.S. (1988). Distribution and magnitude of eagle/livestock conflicts in the western United States. Proc. Vertebr. Pest Conf. 13, 241244. Rabor, D.S. (1965). The present status of the Monkey-eating eagle, Pithecophaga jefferyi Ogilvie-Grant, of the Philippines. Pp. 55-63 in Conference on Conservation of Nature and Natural Resources in Tropical South East Asia, Bangkok, Thailand, November 29 December 4, 1965. International Union for Conservation of Nature and Natural Resources.

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Redig, P.T. (1993). Medical Management of Captive Birds of Prey. Minnesota: The University of Minnesota Press. Reynolds, R.T. (1972). Sexual dimorphism in Accipiter hawks: a new hypothesis. The Condor, 74, 191-197. Rettig, N.L. (1978). Breeding behavior of the Harpy eagle (Harpia harpyja). The Auk, 95(4), 629-643. Rickart, E.A., Heaney, L.R., Heideman, P. D., & Utzurrum, R.C.B. (1993). The distribution and ecology of mammals in Leyte, Biliran and Maripipi Islands, Philippine. Fieldiana:Zoology, N.s. 72. Rodriguez-Estrella, R., Donazar, J.A., & Hiraldo, F.. (1997). Raptors as indicators of environmental change in the scrub habitat of Baja California Sur, Mexico. Conservation Biology. 12(4), 921-925. Rowley, I. (1983). Re-mating in birds, In Bateson, P. [ed.], Mate choice. Cambridge, England: Cambridge Univ. Press. Salvador, J.I.D & J.C. Ibanez. 2006. Ecology and conservation of Philippine Eagles. Ornithological Science. 5, 71-176. Selander, R. K. (1966). Sexual dimorphism and differential in birds. Condor, 68, 113-151. niche utilization

Shiroff, A. (2002). "Paradoxurus hermaphroditus" (On-line), Animal Diversity Web. http://animaldiversity.ummz.umich.edu/site/accounts/information/ Paradoxurus_hermaphroditus.html. [Accessed April 03, 2008]. Tadena, D., Salvador, D.I., Miranda, H.C.Jr., & Ayaay, A.. (2000). Captive breeding by natural pairing, incubation and chick-rearing methods for Philippine eagles. Sylvatrop: Technical Journal of the Philippine Ecosystem Research Development Bureau, DENR. 10 (1 & 2), 59-69 Tigner, J. R., & Larson, G.E. (1981). Golden eagles: scavengers and predators on domestic lambs. In: 1982 Wildlife-Livestock Relationships Symposium. Moscow: Univ. of Idaho. United Nations Environmental Program/Convention on Biological Diversity. (2003). Convention on biological diversity, text and annexes. Canada: Secretariat of the Convention on Biological Diversity. Ward, J.I., Gutierrez, R.J., & Noons, B.R. (1998). Habitat selection by Northern Spotted Owls: the consequences of prey selection and distribution. The Condor, 100, 79-92.

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Whitehead, J. (1899). Field notes on birds collected in the Philippine islands in 1893-96. Ibis. 7(5), 81-111. Wimberger, P. H. (1984). The use of green plant material in bird nests to avoid ectoparasites. Auk. 101, 615-618. Wischusen, E.W. (1990). The foraging ecology and natural history of the Philippine flying lemur (Cynocephalus volans). Ph.D. dissertation, Cornell University (New York: Ithaca). Wischusen, E.W., & Richmond, M.E. (1998). Foraging Ecology of the Philippine Flying Lemur (Cynocephalus volans). Journal of Mammalogy, 79(4), 1288 - 1295. Wood, D. T. (1946). Eye-witness account of golden eagle killing a calf. Condor, 48, 143. Wylie, S.R. (1974). Notes on egg-laying in the Monkey-eating Eagle. Auk, 91, 191. Young, L.S. & Kochert, M.N. (1987). Marking techniques. In Giron-Pendleton, B.A., Millsap, B.A., Cline, K.W., & Bird, D.M. [Eds]. Raptor Management Technique Manual. Washington: Wildlife Federation.

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APPENDIX A Photos of Philippine Eagles, nests and/or nest sites

Fig.1. White lauan Shorea contorta nest tree at Mount Sinaka (red arrow pointing to the nest).

Fig.2. One of two observation blinds used during 1999-2000 observation at Mount Sinaka

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Fig.3. Nest site at Tarragona, Davao Oriental

Fig.4. Female eagle incubating on the Tarragona nest (Jan 2007)

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Fig.5. Nest tree in Mt Apo, Davao City with adult female and one month old chick-(black arrow, April 2006)

Fig.6. Adult female feeding chick with a gravid female Waglers Pit Viper Tropidolaemus wagleri.

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Fig.7. Nesting site at Cabuaya, Mati City.

Fig.8. Adult female with 4-5 month-old chick (June 2005)

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Fig.9. Nest site at Mt Kitanglad, Bukidnon with mature pine trees on foreground.

Fig.10. Three month old chick on the nest doing flight exercises.

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APPENDIX B Data sheet used in studying Philippine Eagle behavior on the nest

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APPENDIX C Philippine Eagle Ethogram 1. General Maintenance Behavior Sleeping SLP One or both eyes are closed. Sometimes, the head is inserted in the feathers of the back. This is done while incubating or at perch. Perched PRD Perched at a branch. Soaring SAR Flying without flapping the wings. Employed by eagles during thermal utilization, the flight pattern is a spiraling ascent. Gliding GLD Flying without flapping the wings but not soaring. Flapping Exercise FEX Powerful and continuous flapping of the wings above the nest but not soaring. Normally done by a juvenile as an exercise. Preening PRN - Using the beak, fixing and arranging the feathers by wiping each feather with the natural oil produced by the preen gland. Sun Bathing SNB Bathing under the sun while both wings are spread open. Usually done during the morning and after a strong rain. Hang HNG Hanging while the wings are open. Normally done by juveniles as an exercise. Defecation DEF As written. Feaking FEK Wiping the beak on a branch after eating. Feeding FED As written. Hunting HUG Flying above the forest, perching and observing the prey. Ends with catching a prey from the ground or from trees.

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Nest-Building NBL Putting fresh or dried twigs or sprigs on the nest. Sprig Delivery SDL Cutting pieces of a branch in nearby trees to bring to the nest. Egg Turning ETN Standing, slowly pushing and placing the egg between the feet using the beak and incubating it.

2. Social Behavior Prey Delivery PDEL The male delivering food for the consumption of the female or the chick. Done by the female for the chick as well. Intruder Chase ICS Flying close to an intruding Philippine Eagle or another raptor, without directly attacking or fighting it. It ends when the intruding bird leaves the territory. Aerial Courtship ACP Mutual flight where pair soars in a circular pattern followed by a rapid dive by the female with the male in hot pursuit Wing Slap WSLAP Strong slapping or flapping of the wings. Allo-grooming AGM The male fixes and arranges the feathers of the female. The female may respond by fixing and arranging the males feathers. Billing BIL The male gently pecks the females beak and female may respond by doing the same. Courtship Feeding CFD The male brings the female food. This is normally followed by copulation. It is coupled with loud calls from the male. Pre-copulation Vocals PCV Loud, short and successive male calls before copulation.

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Copulation COP The male mounts the female and spreads open the cloaca. Loud, short and successive calls would be heard from the male. The female remains silent. Egg lethargy ELG Happens only to the female eagle a few days before egglaying. The female stays at perch or in the nest and is normally sleeping. Some captive eagles normally do not eat for a couple of days during this period. Egg Laying ELN As written. Incubating INC - As written. Brooding BRD The parent eagle will cover the chick with his/her breast feathers. Object Play OPY Repeatedly grabbing a branch, leaf or any other object on the ground or in a tree, coupled with flapping of the wings. Wing Cup WCUP The wings are spread open and the crest are standing. Feeding the Young FYN As written.

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APPENDIX D Sample of transmitter attachment and window clipping method to mark Philippine Eagles

Fig. 1. Transmitter (blue arrow) is harnessed as a backpack to an eagle using a Y-attachment method.

Fig. 2. Sample of a window (black arrow) clipped on three secondary feathers of a wing of a Philippine Eagle.

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APPENDIX E Mean weight used for calculating biomass of prey items and the references. Species Mammals
Cynocephalus volans Paradoxurus hermaphroditus Petinomys crinitus Macaca fascicularis Canis familiaris domesticus Felis domesticus Unidentified mammal Unidentified rodent Unidentified bat Reptiles Naja samarensis Tropidolaemus wagleri Unidentified snakes Birds Sarcops calvus Gallus gallus bankiva Philippine flying lemur Common palm civet Mindanao flying squirrel Long-tailed macaque Domestic dog Domestic cat 1360 3200 225 5000 500 500 250 255 31 Rickart et al (1993) Shiroff (2002) Rickart et al (1993) This study This study This study This study Rickart et al (1993) Rickart et al (1993) This study This study This study Dunning (2000) Del Hoyo et al (1994)

Common Namea

Weight (g)

References

Southeastern Philippine cobra Wagler's pit viper

500 200 200 170 767

Coleto Domestic chicken

Mindanao flying squirrel weight averaged from published weight of three specimens of the similar-sized Philippine tree squirrel Sundasciurus philippinensis from Rickart et al (1993, pg. 47 ) Unidentified rodents weight averaged from published weight of five specimens of the Common Philippine forest rat Rattus everetti from Rickart et al (1993, pg. 51). R. everretti is an abundant native rat in Mindanao forests. Unidentified bat - weight averaged from published weight of six specimens of the Common short-nosed fruit bat Cynopterus brachyotis from Rickart et al (1993, pg. 23). C. brachyotis is an abundant and widespread bat in Mindanao and elsewhere. Southeastern Philippine Cobra actual weight from a specimen in Davao City Waglers pit viper - actual weight from a specimen in Davao City Unidentified snakes we used weight value for Waglers pit viper as a surrogate

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APPENDIX G Calculation of longevity value for Philippine Eagles in the wild state in Mindanao (formula (adapted from Brown, 1966) Longevitye = 2/(N/A [SMy] + age at sexual maturity Where N = no. of fledged young, A=no. of attempts reaching egg-laying, N/A = breeding success SMy = Number of young that reaches sexual maturity (considering 75 % mortality or 25 % survival), in the absence of actual data for Philippine Eagles, values from other large eagles were used (Note: In temperate climates it appears clear from ringing results that 75% or more of the young of large raptors such as Buzzards Buteo buteo and Ospreys Pandion haliaetus die before reaching sexual maturity (Osterlof 1957, Mayaud 1955); and from scanty data this figure seems to apply to eagles Brown 1966. Longevitye = Using values for Philippine Eagle breeding success (1999-2007 data added to Miranda et al 2000 data) = 2/(0.33 young/pair/yr [.25] = 24.2 yrs (add age as immature, 5-6) =29-30 yrs

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APPENDIX H 2008 weights of male and female Philippine Eagles at the Philippine Eagle Center, Davao City. Male Name Magiting Junior Sir Arny Mindanao Tsai Jag Pag-asa Pagkakaisa Bayani Arakan Maginoo Macanudo Gloria Victoria NDY Freedom Geothermica Kantilan Sultan Kagsabua Chick#20 AVE Weight (kg) 4.850 5.550 5.700 5.400 5.200 4.650 6.150 5.350 4.500 5.100 4.400 5.650 5.600 5.200 5.400 4.200 4.350 4.700 5.200 5.113 6.709 1.3 Name Zamboanga Marikit Kahayag Thor Girlie Princess Maasim Ka Brianne BGR Pangarap Kapayapaan Aling Naty Mia Princess of Tupi Dakila Pin-Pin Tinuy-an Female Weight (kg) 6.600 7.400 7.650 6.450 5.800 6.600 7.700 5.700 7.900 8.150 5.950 7.300 5.900 7.150 5.500 5.600

Dimorphism index = Ave female wt./Ave male wt.

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APPENDIX I Mean weight of male and female birds in several moderate to highly dimorphic birds of prey (Source of weight records: Clark 1999).
Weight (g) male Peregrine Falcon Barbary Falcon Gyrfalcon Saker Falcon Lanner Falcon Oriental Hobby Merlin American Kestrel African Black Eagle Eurasian Sparrow Hawk Pallid Harrier birds birds birds, mammals sometimes mammals and birds mainly birds, but also mammals birds and insects birds and mammals insects, birds and mammals mammals, large birds, reptiles mainly birds, occasionally mammals mainly small birds, but also mammals Montagu's Harrier mainly birds, mammals and reptiles Hen Harrier birds or mammals 290-400 (350) 410-600 (520) 1.5 212-300 (260) 260-435 (360) 1.4 235-385 (310) 380-520 (440) 1.4 117-160 (148) 220-350 (260) 1.8 3800 4800 1.3 97-120 (109) 102-150 (123) 1.1 150-190 (175) 180-245 (220) 1.3 131-223 (193) 141-325 (237) 1.2 450-650 (550) 580-750 (650) 350-480 (430) 900-1300 (1100) 665-900 (775) female 750-1150 (950) 550-720 (640) 1300-2100 (1700) 900-1300 (1050) 550-800 (650) 1.2 1.4 1.5 1.5 1.6 Dimorphism index

English Name

Diet

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CURRICULUM VITAE
Jayson Carungay Ibaez
===================================================================

ADDRESS: Present Employer: Philippine Eagle Foundation, VAL Learning Village, Ruby St. Marfori Heights, Davao City Residence: Telefax: E-mail: PERSONAL DATA: Birthday: June 10, 1974 Place of Birth: Tarlac, Tarlac Married, 2 children (Sofia, 8 and Luis, 2) Spouse: Gliceria Balaquit-Ibanez PROFESSIONAL AFFILIATIONS: Board Member, Wildlife Conservation Society of the Philippines Alumni Member, UPLB Biological Society Alumni Member, UP Palaris Confraternity EDUCATION: M.S. B.S. 2008 Ateneo de Davao University Biology (Zoology) 1996 University of the Philippines, Los Baos Biology, Wildlife Ecology major Blk. 16 Lot 6 Pag-Ibig Countryhomes, Catalunan Pequeo, Davao City Philippines. 082 - 224 -3022 falcon2car@yahoo.com

ELIGIBILITY: Career Service Professional

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WORK EXPERIENCES: June 2002 to present. Field Team Coordinator. Field Research Program. Philippine Eagle Foundation. Supervises research on the biology and ecology of the Philippine Eagle.

2002 June to 2004 June. Instructor 5, College of Science and Mathematics. University of the Philippines-Mindanao. Teaches lecture and laboratory classes in life sciences. 1996 July to May 2002. Field Biologist, Field Research Program, Philippine Eagle Foundation, Implements research on eagle breeding behavior, food habits, juvenile behavior and dispersal, habitat use, etc. Also do information campaigns, and socio-economic surveys among communities in the uplands. Volunteer. Dioscoro S. Rabor Wildlife Collection. Birds and Mammals Section. UPLB Museum of Natural History, College, Laguna. Student Assistant. Dioscoro S. Rabor Wildlife Collection. Birds and Mammals Section. UPLB Museum of Natural History, College, Laguna. Field Assistant. Terrestrial Faunal Inventory of the Tamaraw Conservation Project of the UPLB Foundation, Inc. and the Protected Areas and Wildlife Bureau, DENR at Mount Iglit-Baco, Mindoro. Work includes ocular (point and line transect surveys) and capture (mist-nets for birds and bats, cage and snap traps for rodents) surveys of terrestrial vertebrates. Field Assistant. Terrestrial Faunal Inventory of the Luntiang Alyansa ng Bundok Banahaw and UPLB Wildlife Biology Laboratory at Mount Banahaw de Tayabas, Quezon. Work includes ocular (point and line transect surveys) and capture (mist-nets for birds and bats, cage and snap traps for rodents) surveys of terrestrial vertebrates. Field Assistant. Terrestrial Faunal Inventory of Mount Makiling Forest Reserve as part of the SUAN Biodiversity Project by the UPLB Institute of Environmental Science and Management.

1996 Jan to June.

1995 June to Oct

1995 May

1995 April

1995 March

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PUBLICATIONS: - Refereed 2006 MIRANDA, H.C. and J.C. IBAEZ. A modified Bal-Cha Tri to capture Great Philippine Eagles for radio-telemetry. Journal of Raptor Research. 40:235-237. SALVADOR, J.I.D and J.C. IBAEZ. Ecology and conservation of Philippine Eagles. Ornithological Science. 5:171-176. GOMEZ, R.K.S.C., J.C. IBAEZ and S.T. BASTIAN, JR. Diversity and community similarity of Pteropodids and notes on insectivorous bats in the Arakan Valley Conservation Area, Mindanao. Sylvatrop, the Technical Journal of Philippine Ecosystems and Natural Resources. 15 (1 &2): 1-11. CAYUNDA, E.V., J.C. IBAEZ, and S.T. BASTIAN, JR. Roosting behavior and roost site characterization of Peteropus vampyrus in Malagos Watershed, Davao City. Agham Mindanao (2): 61-72. SILVOSA, M., C. CONCEPCION, J.C. IBAEZ and G. IBANEZ. 2004. Bats in the Geothermal Reservation of Mt. Apo Natural Park. Agham Mindanaw (2): 73 82. 2003 J.C. IBAEZ, H.C. MIRANDA, G.B. IBANEZ, D.S. AFAN, and R.S. KENNEDY. Observations on the breeding behavior of a pair of Philippine Eagles in Mount Sinaka, Central Mindanao, Philippines. Wilson Bulletin. 1145(3):333-336. MIRANDA, H.C., JR., D.I. SALVADOR, J.C. IBAEZ, AND G.B. IBAEZ. 2000. Summary of Philippine Eagle reproductive success, 1978-98. J. Raptor Res. 34:37-41. AFAN, D.S., J.C. IBAEZ, G.B. IBANEZ, G. L. BUESER,. K.M. GATIL, and H.C. MIRANDA, Jr. Notes on movement and behavior of post-fledging juvenile Philippine Eagle at Mount Sinaka, Mindanao. Sylvatrop: Technical Journal of the Philippine Ecosystem Research Development Bureau, DENR. 10 (1 & 2):5969. BUESER, G.L., AFAN, D.S., J.C. IBAEZ , G.B. IBANEZ, and K.M. GATIL. Distribution and conservation of Philippine Eagles in Davao Oriental and Davao Province. Sylvatrop: Technical Journal of the Philippine Ecosystem Research Development Bureau, DENR. 8 (1 & 2):114-120.

2006

2005

2004

2000

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IBAEZ, J.C. G.L. BUESER, AFAN, D.S., , G.B. IBANEZ, and K.M. GATIL. Notes on the avifaunal diversity of the highlands of Davao Oriental and Davao Provinces. Sylvatrop: Technical Journal of the Ecosystem Research Development Bureau, DENR. 8(1 & 2):103113. CONSERVATION AND RESEARCH GRANTS APPROVED 2008 Satellite Transmitter Grant. North Star Science and Technology and American Bird Conservancy. Searching for the Haring-Ibon: what is the number and distribution of Philippine Eagles in Leyte and Samar, Philippines? Sea World & Busch Gardens Conservation Fund. 2007 Saving the Haring Ibon : Research, Local Capacity-Building and Education Campaign on the Endangered Philippine Eagle within the Southern Sierra Madre Protected Landscape, Luzon, Philippines. Ocean Park Conservation Fund-Hongkong. Project MAMBOOGOOK: Conservation research on the endangered Philippine Eagle in the Eastern Mindanao Corridor and Central Mindanao Corridor. Critical Ecosystem Partnership Fund. GAYNAWAAN 2004: Conservation of threatened vertebrates at Mount Sinaka, Central Mindanao, Philippines. BP Conservation Programme Bronze Award. Towards the conservation of the Eastern Mindanao Corridor: Biodiversity Archiving and Assessment Project. Critical Ecosystem Partnership Fund. GAYNAWAAN1 Project: Towards the PRESERVATION, REHABILITATION, AND DEVELOPMENT of the Arakan Valley Conservation Area (AVCA). Global Environment Facility-United Nations Development Program-Royal Netherlands Government Small Grants Facility. 2003 Collaborative work plan for the newly formed Alliance for Research and Conservation of the Philippine Eagle. Critical Ecosystem Partnership Fund.

2005

2004

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Development of a biodiversity monitoring project for the eastern Mindanao corridor. Critical Ecosystem Partnership Fund. Conservation and research on the endangered Philippine Eagles at the Northern Sierra Madre National Park, Luzon (NSMNP). World Wildlife Fund Philippines. RESEARCH REPORTS: - non-refereed 2007 Conservation Research on the Endangered Philippine Eagle in Mindanao. A project terminal report submitted to the Marubeni Energy Services. Conservation and research on the Endangered Philippine Eagle of the Northern Sierra Madre Natural Park, Luzon. Project terminal report submitted to World Wildlife Fund Philippines Back to the wild: experimental release of a captive-bred Philippine Eagle at the PNOC Geothermal Reservation, Mount Apo Natural Park, Mindanao. A research report submitted to the Department of Environment and Natural Resources. 2002. Flora and Fauna of Mount Mahuson Range, Cotabato Province: survey results of a participatory resource assessment. Research report submitted to the Foundation for the Philippine Environment. Conservation of biological diversity in the highlands of Davao Oriental and Davao Provinces with a focus on the Philippine Eagle. A technical report submitted to John and Catherine Mac Arthur Foundation. A preliminary survey of small, non-volant mammals of Mount Banahaw de Tayabas, Quezon, Philippines. Unpublished. Undergraduate Thesis. University of the Philippines at Los Baos, College, Laguna.

2005

1997.

1995.

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PAPER PRESENTATIONS 2006 2nd Long-term Ecological Research and DIVERSITAS symposium held in Central Mindanao University, Musuan Bukidnon on October, 2006. Building civil society capacity for biodiversity conservation within the Eastern Mindanao Biodiversity Corridor: An overview of an on-going project. 3rd Asian Raptor Research and Conservation Network Conference held in Kenting Taiwan on October 2003. Conservation status of the endangered Philippine Eagle Pithecophaga jefferyi. Annual Wildlife Conservation Society of the Philippines Symposium-Workshop on Philippine Biodiversity held at Murcia, Negros Occidental on April 2003. Updates on the radio-telemetry of Philippine Eagles in Central Mindanao: 1998-2002. 2000 Wildlife Conservation Society of the Philippines Symposium-Workshop on Philippine Biodiversity held at Cavite City on April 2000. Observations on the breeding behavior of a pair of Philippine Eagles at Mount Sinaka, Mindanao. 1998 Wildlife Conservation Society of the Philippines Symposium-Workshop on Philippine Biodiversity held at Marina Azul Davao City on April 1998. Notes on the avifaunal diversity of the highlands of Davao Oriental and Davao Provinces.

2003

2003

2000

1998

POSTER PRESENTATIONS 2008 9th Student Conference on Conservation Science. University of Cambridge, UK. March 25-28, 2008. Building civil society capacity for biodiversity conservation within the Eastern Mindanao Biodiversity Corridor: An overview of an on-going project. Conservation International Symposium entitled Defying Natures End: the Africa Context. Antananarivo, Madagascar. June 20-24, 2006. Building stakeholder capacity for biodiversity conservation within the Eastern Mindanao Corridor: An overview of an on-going project.

2006

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2003

3rd Asian Raptor Research and Conservation Network Conference held in Kenting Taiwan on October 2003. Updates on the radio-telemetry studies for Philippine Eagles in Mindanao. REFERENCES

Perry S. Ong, Ph.D. Director, Institute of Biology College of Science University of the Philippines Diliman, Quezon City Email: ongperry@yahoo.com Richard T. Watson, Ph.D. Vice-President, Director-International Program The Peregrine Fund 5662 W. Flying Hawk Lane Boise, ID 83709 Email: rwatson@peregrinefund.org

Dennis I. Salvador Executive Director Philippine Eagle Foundation VAL Learning Village Ruby Street, Marfori Heights Davao City, Philippines Tel. 6382-224-3021 Fax. 6382-224-3022 Email: djis@pldtdsl.net