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PHILIPPINE EAGLE Pithecophaga jefferyi BREEDING BIOLOGY, DIET,

BEHAVIOR, NEST CHARACTERISTICS, AND LONGEVITY ESTIMATE IN
MINDANAO ISLAND, PHILIPPINES

Thesis · March 2007

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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. Ibaňez

March 2007
APPROVAL SHEET

ii
 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 organization’s

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.

iii
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 Master’s 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 I’ve inadvertently omitted, thank you.

Finally, this thesis is for happiness, which in my case would mean Bing, Sofia

and Luis.

iv
 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

v
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.

vi
 TABLE OF CONTENTS

Content Page No.

TITLE PAGE i
APPROVAL SHEET ii
ACKNOWLEDGEMENT iii
ABSTRACT v
TABLE OF CONTENTS vii
LIST OF TABLES xi
LIST OF FIGURES xii

CHAPTER
1 INTRODUCTION
Background of the Study 1
Statement of the Problem 4
Significance of the Study 5
Scope and Delimitation of the Study 8
Definition of Terms 9

2 REVIEW OF RELATED LITERATURE AND STUDIES

Value of Conserving the Philippine Eagle 11
Conservation Status of the Philippine 11
Hatching and Fledging Success 12
Mortality 13
Nesting Density and Population Estimate for 14
Mindanao
Entire Philippine Population Estimate and the 16
Luzon Dilemma
Breeding Biology and Ecology of Philippine Eagles 17
Breeding: Monogamy and Site Fidelity 17
Breeding: Eggs, Timing of Breeding, and 19
Incubation

vii
 Nesting Behavior: Courtship and Pre-Egg Laying 20
Nesting Behavior: incubation and brooding 21
Growth and development of young 22
Diet and Feeding Regime 24
Predation on Domestic Animals 25
Nest Sites and Nest Trees 27

3 METHODOLOGY
Description of Study Area 29
Field Data Collection 33
Data Analysis 35

4 PRESENTATION, ANALYSIS AND INTERPRETATION

OF DATA
Nesting Phenology 38
Nest and Nest Tree Characteristics 39
Reproductive Success 43
Behavior of Adults 44
Pre egg-laying period 44
Courtship displays 44
Incubation stage 47
Sprig deliveries 47
Behavior on the nest 48
Prey delivery rate 50
Brooding stage 51
Sprig deliveries 51
Behavior on the nest 51
Prey delivery rate 53
Post-brooding stage 54
Sprig deliveries 54

viii
 Adult behavior on the nest 57
Prey delivery rate 57
Diel pattern of prey delivery 61
Parental care 63
Nestling Development and Behavior 64
Diet and Food Habits 66
Prey species and food habits 66
Philippine Eagle hunting techniques 70
Predation on domestic animals 71
Philippine Eagle diet and food habits at Apo and 73
Kitanglad nests
Size difference between prey items hunted by male 75
and female eagles
Breeding failures as an effect of food stress 77
Estimates for Philippine Eagle adult longevity 78

5 SUMMARY, CONCLUSION, AND RECOMMENDATION

Summary 79
Conclusion 81
Recommendations 82
REFERENCES 84
APPENDICES
A Photos of Philippine Eagles, nests and/or nesting sites. 90
B Data sheet used in studying Philippine Eagle behavior on 95
the nest.
C Philippine Eagle Ethogram. 96
D Sample of transmitter attachment and “window clipping” 99
method to mark Philippine Eagles
E Mean weight used for calculating biomass of prey items 100
and the references.
F Outcomes of breeding attempts by Philippine Eagle pairs 101
in Mindanao from 1977-2007.

ix
 G Calculation of longevity value for Philippine Eagles in the 106
wild state in Mindanao.
H 2008 weights of male and female Philippine Eagles at the 107
Philippine Eagle Center, Davao City.
I Mean weight of male and female birds in several moderate 108
to highly dimorphic birds of prey.
CURRICULUM VITAE 109

x
 LIST OF TABLES

Number Title Page

1 Local land uses within the five Philippine Eagle nesting 30

sites studied from 1999-2007.

2 Nest sites, season and breeding stages studied in 33

Mindanao from 1999-2007.

3 Actual (complete dates) and estimated (month) 39

occurrences of egg laying, hatching and first flight off
nest at four Philippine Eagle nests in Mindanao from
1999-2007.
4 Physical characteristics of Philippine Eagle nests and 40
nest trees on Mindanao Island (all measures were in
meters).
5 Comparison of sex roles for the incubation stage during 49
daytime observations at three Philippine Eagle nests in
Mindanao.

6 Nest attendance, mean trip duration, brooding bout 52

duration and standing bout duration during daytime
observations at Sinaka and Kitanglad nests in Mindanao.

7 Prey delivery rates during the brooding and post- 58

brooding (early and late) stages of the nesting period in
three Philippine Eagle nests in Mindanao.

8 Quantity of Philippine Eagle prey species observed 67

directly from courtship through the nestling stages at four
nest sites in Central and Eastern Mindanao, Philippines,
from 1999 to 2007.

9 Food-niche breadth (FNB), dietary overlap, and 74

estimated mean weights (g) of prey of Philippine Eagle
pairs at Kitanglad and Mt Apo during the nesting season
in Mindanao.

xi
 LIST OF FIGURES

Number Title Page

1 Location of five (5) Philippine Eagle nesting sites studied 31

in Mindanao, Philippines.

2 Sprig delivery rate at Kitanglad and Mt Apo nests during 54

the post-brooding period. 18 = 18-27 days, etc.

3 Times of the day sprig was delivered at Kitanglad and Mt 55

Apo nests during the post-brooding period. 5 = 0500-
0559 hr, etc.

4 Comparison of the mean (n=3) for male, female and 59

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.

5 Comparison of the mean (n=3) for male, female and 60

combined male-female values for average biomass
delivered per day to similar measures for each of three
nests (Cabuaya, Apo, Kitanglad) in Mindanao.

6 Diel pattern of prey delivery at 3 Philippine Eagle nests 61

during the post-brooding period in Mindanao. 6 = 0600–
0759 hrs, etc.

7 Diel pattern of delivery according to weight of prey at 3 62

Philippine Eagle nests during the post-brooding period in
Mindanao. 1 = 1 – 499 g, etc.

8 Percentage of time spent on diurnal activities by the 65

nestling during the post-brooding behavior at the
Cabuaya, Apo and Kitanglad nests in Mindanao.

9 Comparison between the prey mass categories brought 76

by male and female during the post-breeding phase in
three Philippine Eagle nests observed (Cabuaya, Mt
Apo, Kitanglad) in Mindanao. 1= 1–499 g, etc.

xii
 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

world’s 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

 2

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
 3

writing, and the difficulty of doing research in often rugged and rebel-infested

areas (Salvador and Ibaňez, 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.

 4

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 eagle’s 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.

 5

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 world’s 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
 6

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 Eagle’s 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,
 7

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 PEF’s

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 government’s 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 non-

governmental 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.
 8

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.

 9

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. Breeding is successful when the young bird reached post-

fledging stage.

Brooding stage. That stage in the breeding cycle of birds from hatching until

that period when the parents no longer sits on the young bird to regulate

its body temperature.

Congeners. Belonging to the same genus.

Diet. Refers to the species of prey consumed by breeding Philippine Eagles.

Inter-nest distance. The closest distance between active nests of two

monogamous eagle pairs.

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

rears a young that reached sexual maturity.

 10

Nesting phenology. Study of the timing of breeding and its various phases.

Nest characteristics. Refers to measures of eight nest and nest tree

parameters.

Nest site fidelity. 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 parent’s body.

Post-fledging stage. That stage from the time the young bird flew off the nest

until the young disperses away from its parent’s 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

country’s 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

 12

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).

 13

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
 14

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.
 15

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.
 16

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).

 17

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
 18

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
 19

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).

 20

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,

 21

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
 22

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 parent’s 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.

 23

Both parents continue to provide food for the young during the post-

fledging 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 bird’s 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).
 24

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 post-

fledging 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
 25

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
 26

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 Ibaňez, 2006). Eagles feed on palm civets, wild pig, and a young

deer, which closely resemble domestic cats, backyard pigs, and young goats,
 27

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
 28

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 PEF’s 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 10.5–27˚C, 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

 30

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 1999-
2007.

Nest Local land use

Sinaka, Arakan Agroforestry (e.g. shade coffee, fruit tree propagation),
slash and burn farming
Cabuaya, Mati City Slash-and-burn farming, abaca collection, rattan
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

Cabuaya, Mati City. The nest (06o 32.63' N " 126o 1.60’E) 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.

31
 32

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

 33

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 “look-

and-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 Breeding Stage Observation

Observed Observed hours

Sinaka, Arakan 1999-2000 Pre-laying and nest- 703

building, Incubation, early
brooding
Cabuaya, Mati City 2004-2005 Post-brooding 103
Apo, Davao City 2005-2006 Post-brooding 916
Tubaon, Tarragona 2006-2007 Incubation 74
Kitanglad, 2006-2007 Incubation, brooding and 781
Lantapan post-brooding
 34

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).
 35

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
 36

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 Levin’s

(1968) equation: FNB = 1/ΣPij2, 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 Pianka’s Index (O): O =

ΣPijPik/√ Σ Pij 2ΣPik2 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 Brown’s (1966) equation: Longevitye

= 2/(N/A [SMy] + age at sexual maturity), where N/A = breeding success (N = no.
 37

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 chick’s 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.
 39

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 Sinaka Cabuaya Apo Kitanglad

Egg laid Nov 16, 1999 Dec, 2004 Jan, 2006 Nov, 2006

Egg hatched Jan 13, 2000 Feb, 2005 Mar, 2006 Jan 21, 2007

First flight off - Jul, 2005 Aug 2, 2006 Jun 15, 2007
nest

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
 40

contorta, Sinaka), Red Lauan (Shorea negrosensis, Cabuaya ), Tanguile (Shorea

polysperma, Apo and Tarragona), and Igem (Dacrycarpus imbricatus, Kitanglad).

Nest trees were 35–60 m in height (ave. 46.1 ±13.4 m), and nests were 20–46 m

from the ground (Table 4). Nest trees were large, with DBH averaging 1.75 m. All

nests were on a horizontal branch 2–8 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 Altitude NH DBH TH NW NL NB NSBD NDT

Sinaka 950 29.00 0.91 35.40 - - - - -

Apo 630 45.92 2.32 65.63 1.48 2.40 0.52 1.07 2.94
Kitanglad 1434 38.34 2.34 43.75 1.12 2.27 0.65 0.68 1.90
Cabuaya 779 20.70 1.38 37.81 1.90 2.76 1.20 1.80 8.13
Tarragona 898 31.67 1.77 36.88 1.09 1.56 0.44 0.47 1.67
Average 938.20 33.13 1.75 43.89 1.40 2.25 0.70 1.00 3.66
SD 303.30 9.54 0.62 12.56 0.38 0.50 0.34 0.58 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 (630–779 m altitude), while three

in mid-elevation forests (898-1,430 m altitude). All were logged selectively in the

 41

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

study’s average of 33 m fall well within Kennedy’s 27–46 m range. Nest trees

were equally huge with four located in steep slopes—two at the base, one mid-

way and one near a ridge—while 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 1960’s, 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.

 42

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 0.5–1 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).

 43

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 eagle’s 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 eagle’s

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 chick’s 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.

 44

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
 45

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

female’s 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
 46

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 egg-

laying 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,

 47

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 female’s 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
 48

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
 49

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 Sinaka (N=18) Tarragona (N=8) Kitanglada (N=4)

Nest attendance (%) 83 82 78
Female 74 95 100
Male 26 05 0
Mean trip duration (hr)
Female
<1.0 hr 0.15(SD=0.14, n=34) 0.30 (SD=0.20, n=22) 0.50 (SD=0.24, n=13)
>1.0 hr 4.59 (SD=1.72, n=8) 1.08 (SD=0.10, n=3) 1.60 (SD=0.35, n=2)
Male
<1.0 hr 0.09 (SD=0.04, n=4) 0.38 (n=1) b
>1.0 hr 21.04 (SD=3.82, n=5) >64.25 hr* b
Inc. bout duration (hr)
Female 1.43 (SD=0.81, n=70) 1.00 (SD=0.87, n=33) 0.81 (SD=0.39, n=9)
Male 3.14 (SD=1.70, n=11) 1.52 (SD=0.03, n=2) B
th
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
 50

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 male’s 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 = 0.71–0.61 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.

 51

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 7–355 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

 52

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 = 9–54 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 sun’s 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 Sinaka (N=5) Kitanglad (N=9)

Nest attendance (%) 61 50
Mean trip duration (hr)
<1.0 hr 0.17 (SD=0.19, n=12) 0.42 (SD=0.29, n=47)
>1.0 hr 1.65 (SD=0.38, n=4) 1.28 (SD=0.16, n=6)

Brooding bout (hr) 1.34 (SD=0.76, n=14) 1.14 (SD=1.43, n=31)

Standing bout (hr) - 0.35 (SD=0.28, n=17)

N= observation days, SD=Standard Deviation

 53

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.
 54

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.
 55

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.
 56

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 (70–130 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.
 57

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.
 58

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 Early post- Late post- Post brooding

(≤ 28 days) brooding brooding
(29-80 days) (≥ 81days)
No./ Biomass/ No./ Biomass No./ Biomass/ No./ Biomass/
day day (g) day / day (g) day day (g) day day (g)
Sinaka
combined 0.50 269.17 - - - - - -
female - - - - - - - -
male 0.50 269.17 - - - - - -
Cabuaya
combined - - - - 0.69 1803.85 0.69 1803.85
female - - - - 0.38 825.38 0.38 825.38
male - - - - 0.31 978.46 0.31 978.46
Apo
combined - - 0.89 1002.30 0.78 892.25 0.81 924.86
female - - 0.44 399.52 0.41 399.68 0.42 399.63
male - - 0.33 534.63 0.37 492.57 0.36 505.19
unknown 0.11 68.15 0.03 20.04
Kitanglad
combined 0.69 1823.75 0.86 1205.24 0.56 1131.76 0.67 1159.82
female - - 0.48 780.48 0.12 287.79 0.25 475.91
male 0.69 1823.75 0.38 424.76 0.44 843.97 0.42 683.91
 59

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 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.
 60

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.
 61

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 = 0600–0759 hrs,
etc.
 62

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.
 63

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 60–61 days at

the Davao del Sur nest (Gonzales 1968). In captivity, eggs hatch after 56–57

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 104–164 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 re-

nest 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 8–9 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
 64

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
 65

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
 66

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 post-

brooding 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.
 67

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 No. Frequency Biomass %

(%) (g) Biomass
Mammals
Cynocephalus volans 45 30.0 61200 26.2
Paradoxurus
hermaphroditus 21 14.0 67200 28.8
Petinomys crinitus 8 5.3 1800 0.8
Macaca fascicularis 17 11.3 85000 36.5
Cannis familiaris
domesticus 3 2.0 1500 0.6
Felis domesticus 2 1.3 1000 0.4
Unidentified mammal 22 14.7 5500 2.4
Unidentified rodent 13 8.7 3315 1.4
Unidentified bat 6 4.0 186 0.1

Reptiles
Naja samarensis 1 0.7 500 0.2
Tropidolaemus wagleri 3 2.0 600 0.3
Unidentified snakes 3 2.0 600 0.3

Birds
Sarcops calvus 1 0.7 170 0.1
Gallus gallus domesticus 6 4.0 4602 2.0
Total 151 100.0 233173 100.0
Total mammals 137 90.7 226701 97.2
Total birds 7 4.6 4772 2.0
Total reptiles 7 4.6 1700 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 adult’s
 68

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
 69

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 mid-

elevation 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.
 70

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.
 71

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 woman…asked 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
 72

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
 73

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.
 74

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 Kitanglad Apo

Total prey individuals 52 73
Mammal total individuals 52 60
Bird total individuals 7
Reptile total individuals 6
Mean weight of prey (g) 1758.94 1107.23
FNB 3.93 8.31
Dietary overlap 0.61

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,
 75

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’.

 76

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= 1–499 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
 77

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 post-

brooding 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 1.10–1.70

(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

 78

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 Brown’s (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

 80

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
 81

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.
 82

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.

 83

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.
 84

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 90

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
 91

Fig.3. Nest site at Tarragona,

Davao Oriental

Fig.4. Female eagle incubating

on the Tarragona nest (Jan
2007)
 92

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

Wagler’s Pit Viper Tropidolaemus wagleri.
 93

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

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

 94

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.
 95

APPENDIX B

Data sheet used in studying Philippine Eagle behavior on the nest

 96

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.

 97

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 male’s feathers.

Billing BIL – The male gently pecks the female’s 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.
 98

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 egg-

laying. 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.

 99

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.
 100

APPENDIX E

Mean weight used for calculating biomass of prey items and the references.

Species Common Weight References

Namea (g)
Mammals
Cynocephalus volans Philippine flying 1360 Rickart et al
lemur (1993)
Paradoxurus hermaphroditus Common palm 3200 Shiroff (2002)
civet
Petinomys crinitus Mindanao flying 225 Rickart et al
squirrel (1993)
Macaca fascicularis Long-tailed 5000 This study
macaque
Canis familiaris domesticus Domestic dog 500 This study
Felis domesticus Domestic cat 500 This study
Unidentified mammal 250 This study
Unidentified rodent 255 Rickart et al
(1993)
Unidentified bat 31 Rickart et al
(1993)

Reptiles
Naja samarensis Southeastern 500 This study
Philippine cobra
Tropidolaemus wagleri Wagler's pit viper 200 This study
Unidentified snakes 200 This study

Birds
Sarcops calvus Coleto 170 Dunning (2000)
Gallus gallus bankiva Domestic chicken 767 Del Hoyo et al
(1994)
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
Wagler’s pit viper - actual weight from a specimen in Davao City
Unidentified snakes – we used weight value for Wagler’s pit viper as a surrogate
 101

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
 102

APPENDIX H

2008 weights of male and female Philippine Eagles at the Philippine Eagle
Center, Davao City.

Male Female
Name Weight (kg) Name Weight (kg)
Magiting 4.850 Zamboanga 6.600
Junior 5.550 Marikit 7.400
Sir Arny 5.700 Kahayag 7.650
Mindanao 5.400 Thor 6.450
Tsai 5.200 Girlie 5.800
Jag 4.650 Princess Maasim 6.600
Pag-asa 6.150 Ka Brianne 7.700
Pagkakaisa 5.350 BGR 5.700
Bayani 4.500 Pangarap 7.900
Arakan 5.100 Kapayapaan 8.150
Maginoo 4.400 Aling Naty 5.950
Macanudo 5.650 Mia 7.300
Gloria Victoria 5.600 Princess of Tupi 5.900
NDY Freedom 5.200 Dakila 7.150
Geothermica 5.400 Pin-Pin 5.500
Kantilan 4.200 Tinuy-an 5.600
Sultan 4.350
Kagsabua 4.700
Chick#20 5.200

AVE 5.113 6.709

Dimorphism index = Ave female wt./Ave male wt. 1.3
 103

APPENDIX I

Mean weight of male and female birds in several moderate to highly dimorphic
birds of prey (Source of weight records: Clark 1999).

Dimorphism
Weight (g)
English Name Diet index
male female
Peregrine Falcon birds 580-750 (650) 750-1150 (950) 1.5
Barbary Falcon birds 350-480 (430) 550-720 (640) 1.5
Gyrfalcon birds, mammals 900-1300 1300-2100 1.6
sometimes (1100) (1700)
Saker Falcon mammals and 665-900 (775) 900-1300 1.4
birds (1050)
Lanner Falcon mainly birds, but 450-650 (550) 550-800 (650) 1.2
also mammals
Oriental Hobby birds and 131-223 (193) 141-325 (237) 1.2
insects
Merlin birds and 150-190 (175) 180-245 (220) 1.3
mammals
American Kestrel insects, birds 97-120 (109) 102-150 (123) 1.1
and mammals
African Black mammals, large 3800 4800 1.3
Eagle birds, reptiles
Eurasian Sparrow mainly birds, 117-160 (148) 220-350 (260) 1.8
Hawk occasionally
mammals
Pallid Harrier mainly small 235-385 (310) 380-520 (440) 1.4
birds, but also
mammals
Montagu's Harrier mainly birds, 212-300 (260) 260-435 (360) 1.4
mammals and
reptiles
Hen Harrier birds or 290-400 (350) 410-600 (520) 1.5
mammals
 104

CURRICULUM VITAE

Jayson Carungay Ibañez

===================================================================

ADDRESS:

Present Employer: Philippine Eagle Foundation, VAL Learning Village, Ruby St.
Marfori Heights, Davao City

Residence: Blk. 16 Lot 6 Pag-Ibig Countryhomes, Catalunan Pequeño,

Davao City Philippines.

Telefax: 082 - 224 -3022

E-mail: [email protected]

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. 2008 Ateneo de Davao University

Biology (Zoology)

B.S. 1996 University of the Philippines, Los Baños

Biology, Wildlife Ecology major

ELIGIBILITY: Career Service Professional

 105

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.

1996 Jan to June. Volunteer. Dioscoro S. Rabor Wildlife Collection.

Birds and Mammals Section. UPLB Museum of
Natural History, College, Laguna.

1995 June to Oct Student Assistant. Dioscoro S. Rabor Wildlife

Collection. Birds and Mammals Section. UPLB
Museum of Natural History, College, Laguna.

1995 May 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.

1995 April 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.

1995 March 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.
 106

PUBLICATIONS: - Refereed

2006 MIRANDA, H.C. and J.C. IBAŇEZ. A modified Bal-Cha Tri to

capture Great Philippine Eagles for radio-telemetry. Journal of
Raptor Research. 40:235-237.

2006 SALVADOR, J.I.D and J.C. IBAŇEZ. Ecology and conservation of

Philippine Eagles. Ornithological Science. 5:171-176.

2005 GOMEZ, R.K.S.C., J.C. IBAŇEZ 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.

2004 CAYUNDA, E.V., J.C. IBAŇEZ, 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. IBAŇEZ and G. IBANEZ.

2004. Bats in the Geothermal Reservation of Mt. Apo Natural Park.
Agham Mindanaw (2): 73 – 82.

2003 J.C. IBAŇEZ, 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.

2000 MIRANDA, H.C., JR., D.I. SALVADOR, J.C. IBAŇEZ, AND G.B. IBAÑEZ.
2000. Summary of Philippine Eagle reproductive success, 1978-98.
J. Raptor Res. 34:37-41.

AFAN, D.S., J.C. IBAŇEZ, 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):59-
69.

BUESER, G.L., AFAN, D.S., J.C. IBAŇEZ , 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.
 107

IBAŇEZ, 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):103-
113.

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.

2005 Project MAMBOOGOOK: Conservation research on the

endangered Philippine Eagle in the Eastern Mindanao Corridor and
Central Mindanao Corridor. Critical Ecosystem Partnership Fund.

2004 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.
 108

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.

2005 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.

1997. 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.

1995. A preliminary survey of small, non-volant mammals of Mount

Banahaw de Tayabas, Quezon, Philippines. Unpublished.
Undergraduate Thesis. University of the Philippines at Los Baños,
College, Laguna.
 109

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.

2003 3rd Asian Raptor Research and Conservation Network

Conference held in Kenting Taiwan on October 2003.
Conservation status of the endangered Philippine Eagle
Pithecophaga jefferyi.

2003 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 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 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.

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.

2006 Conservation International Symposium entitled “Defying

Nature’s 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.
 110

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. Dennis I. Salvador

Director, Institute of Biology Executive Director
College of Science Philippine Eagle Foundation
University of the Philippines VAL Learning Village
Diliman, Quezon City Ruby Street, Marfori Heights
Email: [email protected] Davao City, Philippines
Tel. 6382-224-3021
Fax. 6382-224-3022
Richard T. Watson, Ph.D. Email: [email protected]
Vice-President, Director-International
Program
The Peregrine Fund
5662 W. Flying Hawk Lane
Boise, ID 83709
Email: [email protected]

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