Tesis Muniz Lopez PDF

BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia harpyja)
EN ECUADOR
Ruth Muñiz López

TESIS DOCTORAL
JUNIO 2016
RUTH MUÑIZ LÓPEZ
BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia

harpyja) EN ECUADOR.
Facultad de Ciencias
Departamento de Ciencias Ambientales y Recursos Naturales
Universidad de Alicante
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Departamento de Ciencias Ambientales y Recursos Naturales
Facultad de Ciencias
BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia harpyja)

EN ECUADOR
Dª. RUTH MUÑIZ LÓPEZ

Licenciada en Biología por la Universidad de Granada.
Tesis presentada para aspirar al grado de

DOCTORA POR LA UNIVERSIDAD DE ALICANTE
Programa de Doctorado: “Ciencias Experimentales y Biosanitarias”
Dirigida por:
DR. VICENTE URIOS MOLINER

Profesor Titular de la Universidad de Alicante
En Alicante, junio de 2016.
Ruth Muñiz López Vicente Urios Moliner
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Churulla, Cunsi Pindo, Kenguiwe, Chorongo wamani, Nasó wiwé... Águila Harpía
Hembra de águila harpía partiendo de su árbol del nido en la Reserva Faunística Cuyabeno.
Foto: P. Oxford/PCAHE-SIMBIOE
A los ojos de Etsá vuela el espíritu del aire en la selva y, de la mano de Arutam,
guía los pasos de los nungkánmayas acogidos por Ikiam
Foto de portada: P. Oxford /PCAHE-SIMBIOE
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ÍNDICE
Página
CAPÍTULO 1 11
1.1. Introducción 13
1.1.2. Antecedentes en la investigación del águila harpía. 14
1.1.3. El águila harpía: la especie. 18
1.1.4. Relación con la cultura y reconocimientos nacionales. 20
1.2. Justificación de la Tesis Doctoral. 21
1.3. Objetivos de la Tesis Doctoral. 22
1.4. Estructura de la Tesis Doctoral. 25
1.5. Área de estudio, resultados generales y discusión general. 26
1.6. Conclusiones generales de la Tesis Doctoral. 32
1.7. Bibliografía. 34
CAPÍTULO 2 45
Movimientos de águilas harpías Harpia harpyja durante los dos
primeros años desde su nacimiento. 47
Movements of Harpy Eagles Harpia harpyja during their first
two years after hatching. 48
Acknowledgements. 55
References. 56
CAPÍTULO 3 61
Dispersión juvenil del águila harpía Harpia harpyja. 63
Harpy Eagle (Harpia harpyja) juvenile dispersion. 64
References. 71
CAPÍTULO 4 77
Ecología trófica del águila harpía (Harpia harpyja) en Ecuador. 79
Trophic ecology of the Harpy Eagle (Harpia harpyja) in
Ecuador. 80
Introduction. 81
Study area. 82
Methodology. 83
Results. 85
Discussion. 88
References. 93
CAPÍTULO 5 105
Hábitat del área de nidificación y estimación de la densidad de
nidos del águila harpía en la Reserva Faunística Cuyabeno,
Ecuador. 107
Harpy Eagle breeding habitat and nesting density estimation in
Cuyabeno Wildlife Reserve, Ecuador. 108
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Introduction. 109
Study area. 112
Methodology. 114
Results. 117
Discussion. 120
Aknowledgements. 130
References. 131
CAPÍTULO 6 149
Mortalidad del águila harpía en Ecuador. 151
Harpy Eagle mortality in Ecuador. 152
Introduction. 153
Study area, materials and methods. 154
Results. 155
Discussion. 155
Aknowledgements. 159
References. 160
AGRADECIMIENTOS 167
ANEXO FOTOGRÁFICO 173
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CAPÍTULO 1
Figura precolombina de águila harpía, 1600 a.C - 600 d.C. Frontera occidental Panamá-Costa Rica.
Foto: Universidad Vanderbilt. TN. EE.UU.
Que las palabras sigan vivas, las ideas no pernocten, las imágenes dancen al son de los
tambores, la música sea sonidos sonoros que lleguen al corazón de Iwia.
Extraído del poema “Como puma herido”

María Clara Sharupi, poeta indígena de la Amazonía ecuatoriana.
Capítulo 1 Tesis Doctoral Ruth Muñiz López
CAPÍTULO 1
1.1. Introducción.
La palabra Neotrópico (del griego neos = “nuevo”) hace referencia a la región tropical
del continente americano. Este término fue acuñado por Peter Martyr dÁnghiera en
1493 poco después del primer viaje de Cristóbal Colón a América (O’Gorman 1972).
Tal y como hoy se define abarca desde la región central de México hasta el sur de
Brasil, incluyendo Centroamérica, las islas del Caribe y la mayor parte de Suramérica
(Schultz 2005). En ella, la precipitación y la temperatura media son generalmente
elevadas (Antonelli & Sanmartín 2011), aunque existe una alta variación entre las
distintas subregiones, por ejemplo, mientras que al oeste de Colombia el nivel de
precipitación es uno de las más altos del mundo con casi 9000 mm/año (Antonelli &
Sanmartín 2011) en la Amazonía la precipitación varía entre 1500 y 3000 mm / año
(Salati & Vose 1984).
Los ecosistemas tropicales son los más diversos del mundo (Wilson 1988) siendo el
Neotrópico la región biogeográfica con mayor diversidad de rapaces en la tierra y el
centro de origen de la familia Falconidae y del género Buteo (Escobar-Acosta et al.
2006). De las aproximadamente 292 especies de aves rapaces diurnas que existen, 222
se encuentran en los trópicos y 96 en el Neotrópico, de los que 18 son endémicas
(Bildstein et al. 1998), habiéndose identificado 67 en Ecuador (Ridgely & Greenfield
2001).
Las aves rapaces son sensibles a la contaminación y a los cambios en la calidad del
hábitat, por lo que pueden desempeñar un papel importante como indicadores
funcionales de la salud ambiental (Leck 1979, Newton 1979, Peakall & Kiff 1988). Para
conservar las rapaces y sus hábitats deben comprenderse sus ciclos biológicos,
requerimientos de hábitat y las condiciones ecológicas y humanas en las que viven de
forma que se logre preservar tanto como sea posible su distribución natural (Stotz et al.
1996).
Tanto el tamaño como la distribución de las poblaciones de aves rapaces pueden
verse limitados por la disponibilidad de lugares adecuados para criar, la existencia de
recursos tróficos, factores climáticos y presiones antrópicas (Newton 1979). En general,
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poseer gran tamaño, baja fecundidad y longevidad son señales que predicen propensión
a la extinción, pero sólo para aquéllas especies que evitan hábitats modificados
(Laurance 1991). Las grandes aves de presa que son relativamente especializadas en
cuanto a su hábitat y dieta generalmente desaparecen cuando se aíslan en hábitats
(Thiollay 1989). Estas aves, después de las típicamente cinegéticas - es decir,
frecuentemente utilizadas para la cacería- son las primeras en desaparecer durante el
proceso de crecimiento de la población humana y el aumento de la explotación de los
bosques lluviosos (Harris 1984).
1.1.2. Antecedentes en la investigación del águila harpía.
Una de las observaciones iniciales sobre esta especie en estado silvestre fue publicada
por Bond (1927) en donde reportó el descubrimiento de un nido de águila harpía en el
noreste del Brasil a aproximadamente 64 km del estado de Pará, en abril de 1926. En el
nido se encontraron dos huevos y se observó el comportamiento de un individuo adulto,
posiblemente la hembra.
Los primeros trabajos acerca de la ecología reproductiva del águila harpía fueron
llevados a cabo en la región Rupununi, en la República de Guyana durante los años 60
(Fowler & Cope 1964). En esta ocasión se descubrieron dos nidos que fueron descritos
y en donde se monitoreó su actividad para estudiar de forma básica los hábitos de las
aves adultas y juveniles, su dieta y las variaciones en su plumaje. Más tarde, en esa
misma región, Brock (1972) realizó observaciones acerca del plumaje de dos águilas
que habían sido capturadas.
Hasta el trabajo de Neil Rettig, realizado entre los años 1974 y 1975 (Rettig 1978),
apenas se conocía el comportamiento reproductivo del águila harpía. Éste regresó a uno
de los nidos encontrados por Fowler y Cope en el año 1960, y describió el lugar de
nidificación, la construcción del nido, el período de cópula, la puesta de huevos, la
incubación, la alimentación de adultos y cría, los cuidados parentales y el
comportamiento de la cría en las distintas etapas de desarrollo. Más tarde, Izor (1985)
realizó un estudio de las presas aportadas a un nido también en la República de Guyana
y posteriormente, entre los años 1987 y 1988 se describió por primera vez la anidación
de esta especie en la Provincia de Misiones (Argentina) (Chébez et al. 1990). Aún así,
hasta la tesis doctoral realizada por Álvarez-Cordero en la región Guayana de
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Venezuela y en la provincia de Darién en Panamá (Álvarez-Cordero 1996) no se

presenta información detallada acerca de esta especie, incluyendo su rango ecológico y
geográfico, biología de la anidación, independencia del área natal, dieta, y amenazas, así
como propuestas para apoyar su conservación.
Tras el estudio de Álvarez-Cordero, en Panamá se realizaron algunas
investigaciones complementarias acerca de su biología reproductiva, distribución
histórica (Mosquera et al. 2002, Aparicio 2003) y tamaño de población (Vargas &
Vargas 2011).
En Perú los trabajos sobre esta águila son casi inexistentes, excepto por aquéllos
realizados por Piana (2007) acerca de la nidificación, distribución y abundancia del
águila harpía en la provincia amazónica Madre de Dios.
En Brasil existen trabajos acerca de su comportamiento durante el periodo de cría
(Sanaiotti 2002) y hábitos alimenticios (Aguiar-Silva 2014).
Otros datos puntuales se pueden encontrar en la bibliografía acerca de su
comportamiento y ecología del aprovisionamiento alimenticio a lo largo de su rango de
distribución (Eason 1989, Peres 1990, Touchton et al. 2002, Rotenberg et al. 2012) o de
especímenes reintroducidos (Muela & Curti 2005). Por otra parte, es interesante señalar
la existencia de estudios genéticos para esta especie, en los que se describe su cariotipo
(Felipe et al. 2001, De Oliveira et al. 2005), se muestra el origen no monofilético del
grupo Harpiinae (Lerner & Mindell 2005) y se concluye que existen niveles
moderadamente altos de diversidad genética entre subgrupos de águila harpía a lo largo
de su distribución (Lerner et al. 2009)
 En Ecuador:
La primera vez que se recogieron datos de campo acerca de la distribución del

águila de manera formal fue en los años cuarenta, encontrándola al oeste de Ecuador
(Orcés 1944).
En 1990 se afirma que el águila harpía se distribuye en el bosque húmedo tropical
bajo los 600 m s. n. m., tanto en la región occidental (costera) como en la oriental
(región amazónica) (Ortiz-Crespo et al. 1990) del país.
Álvarez-Cordero (1996) recogió en su tesis datos de museos que poseían pieles de
águilas ecuatorianas que databan de 1880 (en el British Museum of Natural History,
Reino Unido), 1925 y 1926 (en el American Museum of Natural History, EE. UU.). Los
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ejemplares procedían de la localidad de Balzar (provincia de Guayas), de la

desembocadura del río Cotapino (suroccidente de la provincia de Orellana) y la
desembocadura del río Curaray (provincia de Pastaza) respectivamente. Los colectores
o referentes para estos restos fueron, siguiendo el orden cronológico, Illingwroth y
Olalla e hijos.
Mauricio Guerrero realizó su tesis de licenciatura con el águila harpía en la
Universidad Central de Ecuador (Quito). Los objetivos fueron realizar una evaluación
del estado poblacional de esta especie en el país y determinar el valor cultural o mítico
que le asignan al águila diferentes grupos humanos en Ecuador (Guerrero 1997). Se
trató de un estudio preliminar en el que tomó algunos datos de ejemplares capturados o
tiroteados, profundizando su enfoque en el ámbito etnozoológico. Según ese trabajo, la
distribución del águila harpía al occidente ecuatoriano es muy restringida, limitándose
actualmente al noroccidente, en la provincia de Esmeraldas, en la región que comprende
la parte noroeste de la Reserva Ecológica Cotacachi-Cayapas, en donde obtuvo un
registro visual en abril de 1991. Concluía que en el oriente o región amazónica
ecuatoriana su distribución es más amplia, coincidiendo con bosques poco perturbados
y escasa presencia humana.
Guerrero registró 16 individuos entre 1989 y 1995, gran parte de ellos en los ríos
Cuyabeno, Zábalo y Pacayacu, afluentes del río Aguarico, en la provincia de
Sucumbíos, nororiente de Ecuador. Otros registros se localizaron en la provincia situada
al sur, Orellana, en los alrededores de los ríos Yasuní y Cononaco, lagunas de
Garzacocha y Jatuncocha, así como en las riberas del río Tiputini. Llegó a localizar tres
nidos, todos en el oriente ecuatoriano, pero ninguno activo en ese momento. En su tesis,
analizó el interés que el águila harpía provocaba en el grupo étnico huaorani (propio de
la provincia de Orellana) en donde coronas y brazaletes de guerra son generalmente
adornados con plumas de esta águila con el objeto de lograr la protección y fuerza que
consideran ofrece esta ave. Literalmente cita: “Ellos la admiran por su fuerza y su
destreza cazadora, deseando imitarla para el mismo fin. La consideran un símbolo
protector de sus comunidades y niños, por lo que las capturan para mantenerlas junto a
ellos; eso significa adquirir un estatus superior y una posición privilegiada, pues
consideran que en el ser del cazador ha sido encarnada toda la sabiduría y habilidad del
águila”.
En la región occidental, en donde los bosques ya han sido fragmentados, el águila
harpía parece estar prácticamente desaparecida (Ridgely & Greenfield 2001). Leck
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(1979) ya hizo referencia a la prácticamente nula existencia de registros de la especie en

esta región, y sugería que probablemente estuvieran sufriendo extinciones locales.
En el año 2001 se publicó el libro “Aves del Ecuador” (Ridgely & Greenfield 2001)
en el que se señala que esta especie se encuentra relativamente extendida al este del
país, en donde todavía permanecen bosques de forma extensiva y añadiendo que, a
pesar de que su territorio sea considerable, los individuos observados siempre son muy
pocos. Según el mismo libro, se ha encontrado por debajo de los 400 m s.n.m. (salvo en
un registro en Tinalandia, un hotel a unos 700 – 800 m s.n.m. al oeste de la cordillera
andina). Al noreste casi todos sus registros son de aves disparadas o que han sido
capturadas para mantenerlas en cautividad a pesar de que está penado por ley (salvo en
casos de tradición cultural ancestral como el de la etnia huaorani, en el oriente
ecuatoriano). En la región occidental sólo obtuvieron un único registro de dos aves en
1983, cerca de Tinalandia.
Más tarde, ya en el año 2004, se encontró el primer y único nido activo de esta
especie localizado hasta el momento al oeste de la cordillera andina (Muñiz-López
2007).
El águila harpía no es considerada como una especie abundante, aún en áreas sin
habitar (Ridgely & Greenfield 2001). En comparación con la región occidental, se
encuentran de forma más regular en la región del bajo río Aguarico (provincia
Sucumbíos, nororiente de Ecuador), en donde aún así existen escasísimos avistamientos
(Ridgely & Greenfield 2001).
La tenencia de esta ave en cautividad se describe desde 1971, en donde Stummer
relata en un artículo la captura de un águila harpía en el occidente ecuatoriano y su
posterior traslado a Seattle, EE. UU. (Stummer 1971). Otras informaciones sugieren la
captura y sacrificio de un ejemplar de esta especie en la comunidad afroecuatoriana de
Mataje (provincia de Esmeraldas, noroccidente de Ecuador), en el área del río Mataje, a
principios de 1991 (Jaime Cevallos, en Guerrero 1997). Además, se determinó que un
águila cautiva con plumaje de juvenil, que se encontraba en una residencia privada cerca
de la Reserva Manglares-Churute (provincia de Guayas, suroccidente del país), murió
en 1995 a causa del ataque de otra harpía cautiva adulta al intentar reproducirlas en
cautividad (Mireya del Pozo, en Guerrero 1997).
La comercialización fue evidente en un local privado en la provincia de Esmeraldas.
Allí habían mantenido tres ejemplares de águila harpía en diferentes años. De los tres
individuos, uno había estado cautivo en las inmediaciones de la Reserva Manglares-
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Churute, y se desconoce su destino; el segundo individuo formaba parte de la colección

del zoológico de Salango (provincia de Manabí, occidente de Ecuador), aunque después
fue integrado al programa de reproducción en cautividad de esta especie desarrollado
por la institución “The Peregrine Fund”, regresando a un centro de fauna privado de
Ecuador al finalizar el mismo, en el año 2006 (The Peregrine Fund 2006). Del tercer
ejemplar no se tiene información exacta, ya que se declaró que desapareció en 1991, no
descartándose su posible comercialización (Guerrero 1997).
En el año 2007 fue localizada otra águila harpía de unos dos meses de edad que se
encontraba mantenida en cautividad en una casa de la comunidad San Pablo de
Katetsiayá, perteneciente a la nacionalidad indígena Secoya o Siecopai, en la región
amazónica ecuatoriana. Según la persona que la había capturado, derribó el árbol en
donde se encontraba el nido para recoger al pollo y vendérselo a un traficante que meses
antes pasó por el lugar ofreciéndose para comprar una. Sin embargo, de acuerdo con el
captor y al contar el ejemplar con poca edad, se reinsertó con éxito de nuevo al bosque,
en donde la pareja adulta de águilas silvestres continuó cuidado de este pollo hasta al
menos un año después de su liberación (Muñiz-López et al. 2008).
El primer nido activo de águila harpía que fue monitoreado en Ecuador fue
encontrado en el año 2002, al noreste del país (Muñiz-López 2007). Desde entonces,
quince nidos más han sido localizados en la región amazónica ecuatoriana y uno en la
región occidental de este país.
1.1.3. El águila harpía: la especie.
El águila harpía (Harpia harpyja) es el accipítrido de mayor tamaño del continente

americano (Collar 1989) y una de las cuatro águilas más grandes del mundo, junto con
el pigargo de Steller del este de Siberia (Haliaeetus pelagicus), el águila marcial de las
sabanas africanas (Polemaetus bellicosus) y el águila pitecófaga (Pithecophaga jefferyi)
que habita en cuatro de las islas de Filipinas (Collar 1989, Sick 1997). De todas ellas, es
la que posee las garras de mayor longitud del mundo (Brown & Amadon 1968), estando
considerada como la más poderosa de todas las de su grupo (Collar 1989).
El águila harpía se distribuye a lo largo de bosques tropicales y subtropicales, en
general por debajo de los 900 msnm (Stotz et al. 1996) desde el sur de México hasta el
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norte de Argentina (Hilty & Brown 1986), aunque la población desde México hasta
Costa Rica se encuentra muy debilitada (Vargas et al. 2006).
En Ecuador su presencia se encuentra restringida a algunos parches de bosque al
noroeste del país, pero más homogéneamente en la región amazónica (Guerrero 1997,
Ridgely & Greenfield 2001, Muñiz-López 2003) en donde la temperatura oscila
alrededor de los 25º C y las precipitaciones se calculan entre los 2.000 a 4.000 mm
anuales, siendo el promedio de humedad ambiental entre un 96-100%. (Sierra et al.
1999).
El águila harpía cría un polluelo cada dos años y medio o tres años y se presume
que permanece en el territorio de los padres durante al menos dos años antes de
independizarse del área paterna (Rettig 1978, Ruschi 1979, Álvarez-Cordero 1996).
Su alimentación se basa fundamentalmente en mamíferos arborícolas,
complementándose por aves y algunos reptiles (Fowler & Cope 1964, Rettig 1978,
Álvarez-Cordero 1996, Muñiz-López et al. 2007).
Estas águilas son especialmente longevas. En la República de Guyana se estima que
una pareja utilizó el mismo nido durante más de 30 años (Fowler & Cope 1964).
El águila harpía es una rapaz forestal. Raras veces se le ha visto planeando por
encima del bosque (del Hoyo et al. 1994). En vuelo las alas se notan cortas en
proporción con el resto del cuerpo (180 a 200 cm), así como muy anchas y redondeadas
(del Hoyo et al. 1994).
Poseen un disco facial que puede aumentar su capacidad auditiva y ojos muy
frontales que son útiles para localizar con facilidad sus presas y medir las distancias a
éstas con efectividad, pues se mejora su visión tridimensional (Ferguson-Lees &
Christie 2001). No adquieren su plumaje definitivo hasta los cinco años de edad
(Ferguson-Lees & Christie 2001).
Como en muchas otras rapaces, la hembra es de mayor tamaño y peso que el macho
(del Hoyo et al. 1994). La primera puede pesar hasta 9 kg (6,0 – 9 kg), a diferencia del
macho que llega como mucho a los 5 kg (4 – 5 kg) (del Hoyo et al. 1994). La longitud
de su cuerpo oscila entre los 89 a 105 cm (del Hoyo et al. 1994).
Su gran tamaño hace difícil la confusión con otras especies si se la ha visto alguna
vez. Sin embargo, a veces pueden existir problemas para discernirla del águila crestada
(Morphnus guianensis), aunque esta última es más pequeña, posee una cresta sin
bifurcar y sus tarsos resultan mucho más finos a simple vista (del Hoyo et al. 1994).
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Actualmente, se cataloga de forma global como “Casi Amenazada” en todo su

rango (Collar et al. 1994, BirdLife International 2013) pero en Ecuador es considerada
“Vulnerable” (Granizo et al. 1997), aunque en la región occidental del país podría
incluirse en la categoría de “Peligro Crítico” (Ridgely & Greenfield 2001).
1.1.4. Relación con la cultura y reconocimientos nacionales.
La relevancia a nivel cultural del águila harpía es, en general, un campo poco conocido.
Sin embargo, está considerada como una “Especie Integral” (especie con valor para la
biodiversidad y para la cultura) (Muñiz-López 2002).
Se han reportado registros de tallas, representaciones y figuras en la cultura Olmeca
de México (1500-500 d. C.) (Pool 2007), en la cultura Veraguas de Panamá (500-1200
a. C.) (Newton 1987), en la cultura Tairona de Colombia (1000-1400 a. C) (Mason
1931), en la cultura Saladoide-Barrancoide (ver Figura 1) de Venezuela (500-280 a. C.)
(Silverman & Isbell 2008), en las culturas Tolita (400 a. C.) (DeBoer 1996) y Chorrera
(1200 a. C.-500 d. C.) (Gutiérrez-Usillos 2002) de Ecuador y en la cultura Chavín de
Perú (900-200 d. C.) (Fagan 1996).
Figura 1: Cerámica barrancoide del Bajo Orinoco. Cabeza dimófica superpuesta con un águila
harpía (Silvermand and Isbell 2008)
En las culturas indígenas amazónicas, el águila harpía representa al espíritu del aire
y es el alter ego de los shamanes en el cielo, tal y como el jaguar lo es en la tierra y la
anaconda en el agua (Beltrán 2013). Los shamanes son los líderes intelectuales de las
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culturas nativas americanas. En sus trances se transforman en águila harpía para que
éstas sean las mensajeras desde el mundo espiritual al terrestre. Entre otras muchas
habilidades, pueden negociar con los dueños del mundo animal a través de su espíritu
para lograr que la cacería o pesca sea abundante (Muñiz obs. pers). Un shamán puede
ser iniciado por el águila harpía, adquiriendo la capacidad de volar, que a su vez le
brinda una cosmovisión del universo; sus plumas les conferirán fuerza y poder (Reina &
Kensinger 1991).
Algunos países han dado un reconocimiento especial en su patrimonio al águila
harpía. El 22 de Agosto de 1996 fue declarada Monumento Natural de Argentina de
acuerdo a la Ley Nº 2932 de Áreas Naturales Protegidas y de Interés Público.
Posteriormente, fue nombrada Ave Nacional de la República de Panamá, amparada por
la Ley 18 del 10 de Abril de 2002. Ese mismo año, en el mes de julio, el águila harpía
pasaba a ser el “Ave Representativa de la Diversidad Biológica del Ecuador”, mediante
el Acuerdo Ministerial número 089, y Registro Oficial número 635.
Tras esto, en Ecuador se escribe la “Estrategia Nacional para la Conservación del
Águila Harpía” que, validada y reconocida a través de un Acuerdo Ministerial en marzo
de 2007, y tras ser publicada en el Registro Oficial número 24 un año después, forma
actualmente parte de la legislación ambiental del país (Muñiz-López et al. 2008).
1.2. Justificación de la Tesis Doctoral.
Suramérica contiene aproximadamente 92 especies de aves rapaces diurnas, siendo

Ecuador uno de los cuatro países con mayor diversidad del continente americano (67
especies) (Chébez 2001). A pesar de ello, todavía se desconoce la historia natural,
distribución, abundancia, ecología y necesidades de muchas de ellas, lo que dificulta el
diseño de acciones útiles para su conservación. Existe la necesidad urgente de
documentar estas especies desde diferentes puntos de vista, particularmente en los
trópicos en donde se encuentra la mayor parte de aves rapaces amenazadas y en donde
la destrucción de hábitat ocurre a mayor velocidad (Bildstein et al. 1998)
El águila harpía es una especie de costumbres reservadas que habita bosques
húmedos de zonas remotas de parte de Centroamérica y Suramérica, y por ello es difícil
de detectar (Álvarez-Cordero 1996). Obtener los suficientes datos o un tamaño de
muestra adecuado para estudiarla es particularmente complicado y requiere una fuerte
21
inversión de tiempo (Bednarz 2007), lo que ha resultado en un escaso número de

investigaciones que la hayan abordado a pesar de su amplia distribución (Vargas et al.
2006). Dentro del grupo de los accipítridos, existen nueve especies catalogadas como
“Críticamente Amenazadas”, ocho como “Amenazadas” y 26 como “Vulnerables”,
entre las que se encuentra el águila harpía (BirdLife 2015). La mayoría de ellas se ven
seriamente afectadas por la destrucción de hábitat y la persecución humana (Remsen et
al. 2011).
Antes de crear e implementar un plan de manejo o conservación de cualquier
especie, ésta debe ser estudiada y comprendida. Su estudio es el primer paso para su
protección. Con esta tesis se documentan aspectos de la historia natural, características
que definen su vulnerabilidad y necesidades de conservación de un ave carismática pero
apenas conocida en Ecuador, aportando información que puede ser utilizada para
recomendar acciones que favorezcan su protección. En base a lo anterior, la carencia de
conocimiento biológico de esta especie en el país justifica la elaboración del presente
estudio.
1.3. Objetivos de la Tesis Doctoral.
Esta Tesis Doctoral tiene como objetivo principal contribuir al conocimiento del águila
harpía proporcionando información biológica que caracterice a esta especie en Ecuador
y aportando propuestas para la gestión y conservación de sus poblaciones (Capítulos 2,
3, 4, 5 y 6).
Objetivos específicos:
1.3.1. Caracterizar la dinámica de movimiento de individuos juveniles y establecer

su periodo de dependencia del área natal. Investigar la hipótesis de que la
dependencia juvenil supera el año desde el nacimiento del pollo. Se plantean
las siguientes preguntas (Capítulos 2 y 3):
- ¿Cuánto tiempo permanecen los juveniles en el territorio paterno?

- ¿Cuánta distancia recorren los juveniles durante sus primeros dos años
de vida?
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- ¿El distanciamiento se produce de forma progresiva o repentina?

- ¿La dispersión de los distintos individuos ocurre de forma similar?
- ¿Qué recomendaciones surgen para la conservación de esta especie?
1.3.2. Determinar el comportamiento de dispersión juvenil. Contrastar la hipótesis

de que a partir de los dos años y medio de edad los juveniles abandonan el
área paterna para comenzar su dispersión (Capítulo 3).
 ¿A qué edad comienzan los juveniles sus movimientos de dispersión?

 ¿Qué distancia alcanzan desde su área de cría?
 ¿Qué área llegan a ocupar mientras se dispersan?
 ¿Existe algún lugar de asentamiento en el área que exploran los juveniles
durante su primera etapa de dispersión?
1.3.3. Describir la dieta y ecología trófica durante el periodo de cría en distintas

áreas de nidificación. Testar la hipótesis de que la mayor contribución a la
dieta del águila harpía proviene de mamíferos arborícolas (Capítulo 4).
- ¿Cuál es el espectro de especies que caza el águila harpía cuando está

criando y qué características tienen?
- ¿Es el águila harpía especialista en cuanto a su alimentación?
- ¿Cuándo y por quién se producen los aportes de alimentación al pollo?
¿Varían según las distintas parejas y del estado de desarrollo en el que se
encuentren los pollos?
- ¿Qué preferencias de caza poseen los parentales?
1.3.4. Estimar la densidad de nidos en una región del nororiente de Ecuador

describiendo las características del paisaje en donde el águila harpía
establece su área de nidificación, de lo que se genera un mapa de
distribución potencial en Ecuador. (Capítulo 5).
- ¿Cuántas parejas de águila harpía puede contener un área protegida como

la Reserva de Cuyabeno en Ecuador?
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- ¿Cuáles son las características del paisaje en donde las águilas harpías
establecen sus áreas de cría en el área de estudio? ¿Qué elementos
influencian más la presencia de nidos?
- ¿En qué áreas de Ecuador podrían tener un hábitat adecuado para criar?
¿De cuántas hectáreas disponen?
1.3.5. Examinar las causas de mortalidad en el norte de la región amazónica de

Ecuador (Capítulo 6).
- ¿Cuál es es la tasa de mortalidad detectada en el área de estudio para

adultos y juveniles?
- ¿Existe algún momento crítico en el desarrollo de los juveniles que
pueda hacerlos más sensibles a fallecer?
- ¿Qué condiciones y razones originaron la muerte de los ejemplares
localizados?
- ¿Cuáles pueden ser las acciones para disminuir la mortalidad de esta
especie?
1.3.6. Identificar los factores de amenaza que pueden afectar al águila harpía a lo
largo de su rango de distribución en Ecuador y proponer lineamientos que
puedan contribuir a su conservación (Capítulos 2, 3, 4, 5 y 6).
- ¿Qué acciones pueden emprenderse para reducir el impacto y grado de

presión antrópica hacia el águila harpía?
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1.4. Estructura de la Tesis Doctoral:
La presente memoria doctoral se compone de un capítulo introductorio y de

resumen de memoria de tesis y cinco capítulos estructurados en formato de publicación
científica. De forma adicional se incluye un anexo fotográfico para ilustrar el trabajo de
campo realizado.
A este capítulo (Capítulo 1), le siguen los Capítulos 2, 3, 4 ,5 y 6 que reproducen

el contenido de artículos publicados, en revisión o encaminados hacia su publicación en
diferentes revistas científicas, por lo que se presentan en inglés, con sus
correspondientes secciones de introducción, material y métodos, resultados y discusión.
Al inicio de cada capítulo se presenta el resumen del artículo traducido al castellano.
Cada capítulo tiene su propia sección de referencias y el formato de las citas puede no
ser concordante entre ellos.
A continuación se describe el contenido de los siguientes capítulos:
En el Capítulo 2 (Muñiz-López, R., R. Limiñana, G.D. Cortés and V. Urios. 2012.

Movements of Harpy Eagles Harpya harpija during their first two years after hatching.
Bird Study 59 (3): 1-6) se describen por primera vez los movimientos águilas harpías
silvestres durante sus primeros dos años de vida. Para ello se capturaron dos individuos
y se equiparon con un transmisor de satélite con GPS y panel solar, obteniéndose
posiciones geográficas para cada uno de ellos. Se evaluó cómo ocurría el
distanciamiento al árbol del nido con el paso del tiempo y se compararon los dos casos
para determinar diferencias en esta tendencia.
El Capítulo 3 (Urios V., R. Muñiz-López and J. Vidal-Mateo. Harpy Eagle Harpia
harpyja juvenile dispersion) complementa la información obtenida para los individuos
equipados con emisores GPS del capítulo 2, analizándose cuándo ocurre el momento de
dispersión juvenil y de qué manera progresa su distanciamiento del área natal. Además,
se estudian las distancias horarias antes y después del comienzo de la dispersión y el
área que ocuparon los juveniles durante su periodo de dispersión mientras el emisor se
encontró en funcionamiento.
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A lo largo del Capítulo 4 (Muñiz-López R. Trophic ecology of the Harpy Eagle

(Harpia harpyja) in Ecuador) se examinan los hábitos y preferencias alimenticias del
águila harpía durante su periodo de cría, analizando su grado de especialización y la
estrategia trófica que adopta en diferentes áreas de nidificación. Además se estudia la
contribución de los parentales a la alimentación del juvenil a lo largo de su desarrollo y
las preferencias de caza de los adultos según los hábitos de sus presas.
En el Capítulo 5 (Muñiz-López R. & H. Alcántara. Harpy Eagle breeding habitat
and nesting density estimation in Cuyabeno Wildlife Reserve, Ecuador) se analizan las
características a nivel de paisaje de las áreas de cría de águila harpía en la región
nororiental de Ecuador y se realiza una estimación de la densidad de nidos que puede
existir en el área estudiada. Además, se genera un mapa que perfila la distribución
potencial a nivel de país y se examina el grado de alteración por causas antrópicas que
existe en las áreas en donde parejas de águila harpía pudieran estar criando. Asimismo
se sugieren algunas acciones que pueden tomarse en cuenta para mejorar o mantener las
condiciones idóneas de su hábitat.
En el Capítulo 6 (Muñiz-López R. Harpy Eagle mortality in Ecuador) se analizan
las causas de muerte registradas durante el periodo de estudio. Se recopilan los casos de
individuos fallecidos tanto por muerte natural como por razones antropogénicas y se
examinan los motivos que pudieron dar origen a cada una de ellas, señalando algunas
recomendaciones para disminuir su mortalidad.
1.5. Área de estudio, resultados generales y discusión general.
La presente tesis recoge aspectos generales pero inéditos de la biología del águila de
mayor tamaño del continente americano, el águila harpía (Harpia harpyja), realizando
un recorrido por aspectos como su dispersión juvenil- desconocido en el mundo hasta la
presente investigación-, ecología trófica, hábitat, densidad poblacional, mortalidad y
recomendaciones para su conservación.
El estudio fue realizado en Ecuador (Mapa 1), principalmente en la Reserva
Faunística Cuyabeno y su área de amortiguamiento, situada en la región amazónica al
nororiente del país, y en un área dentro del territorio afroecuatoriano al noroccidente
(Mapa 2).
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Mapa 1: Localización geográfica de Ecuador (área de color marrón) en el continente americano.
La Reserva Cuyabeno, localizada entre las provincias de Sucumbíos y Orellana, es

fronteriza con Colombia y Perú, y se encuentra en la región amazónica ecuatoriana
(Sierra et al. 1999), a una altitud entre 250 y 326 m s. n. m. (Cañadas 1983), con una
temperatura media entre 24,6 y 26,3 °C y una precipitación media de entre 2093,8 y
2728,2 mm al año (Cañadas 1983). La riqueza biológica de esta reserva es
extraordinaria, contando con la mayor densidad de especies arbóreas del mundo
(Valencia et al. 1994). Cinco nacionalidades indígenas reparten sus territorios en esta
área, conformando una amalgama cultural que caracteriza socialmente a la reserva
(ICCA 2010).
En la región occidental, el área de estudio se concentró en la provincia de
Esmeraldas, que forma parte de la región del Chocó ecuatoriano en donde existen
bosques tropicales húmedos y muy húmedos (Sierra et al. 1999). La humedad relativa
media supera el 90%, la temperatura media varía de 23 a 25 °C y la pluviosidad media
anual comprende de 3.500 a 4.000 mm (Jahn 2011). Esta área contiene una rica
biodiversidad, calculándose unas 830 especies de aves y unas 6.000 especies de plantas
(Critical Ecosystem Partnership Fund 2001). Los habitantes de esta región corresponden
a la cultura afroecuatoriana (Minda-Batallas 2002).
27
Mapa 2: Coloreadas de verde se muestra la localización del área de estudio en la Reserva Faunística
Cuyabeno al oriente y territorio afroecuatoriano al occidente de Ecuador
La información de esta tesis proviene de los datos recogidos en las áreas de cría de
águila harpía, localizados en colaboración con las comunidades locales y siguiendo sus
rutas de cacería o pesca. Próximo a algunos de los nidos se armaron torres de
observación de aproximadamente 28 – 30 m de altura para mejorar las condiciones de
visibilidad, puesto que las plataformas de los nidos son construidas en árboles
emergentes como ceibos (Ceiba pentandra) o cedrelingas (Cedrelinga cateniformis).
Para obtener los datos de dispersión juvenil, dos águilas fueron capturadas y equipadas
con emisores satélite con GPS de 70 g. El seguimiento se realizó hasta que los juveniles
contaron con 27 y 39 meses de edad.
Según nuestros resultados, la dispersión juvenil del águila harpía no comienza hasta
el mes 28 de edad, siendo uno de los periodos más largos de dependencia de los
parentales que se conoce para aves rapaces. Su área de dispersión, que se calculó en 386
km2, y las distancias recorridas, 1,07 km/día como media llegando como máximo a
35,1 km del nido con 39 meses de edad, son menores que las observadas para otras
especies de grandes águilas, probablemente debido a las condiciones de homogeneidad
del hábitat que les ofrece el bosque tropical en el que se encuentran. Uno de los
juveniles seguido a través del emisor satélite y que no se desplazó de su área de cría a
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pesar de haber llegado el momento en el que era esperable que comenzase su dispersión,
murió tras haberse observado que semanas antes era atacado por uno de los adultos. Es
la primera vez que se advierte este comportamiento en el águila harpía, aunque ya ha
sido reportado para otras especies, ya que la agresión hacia los juveniles favorece la
independencia y el comienzo de la dispersión.
A pesar de que el águila harpía puede depredar sobre una variedad de especies,
mayoritariamente mamíferos arborícolas, existe especialización en su dieta, prefiriendo
monos (diversas especies) y perezosos (Choloepus spp. y Bradypus variegatus) como
parte de su alimento. Las águilas harpías capturan preferiblemente animales que utilizan
habitualmente el estrato más alto del bosque, y son los machos los que alimentan a su
cría de forma más frecuente, entregando el alimento en la primera fracción del día y
cada 3,8 días. Aunque en todos los nidos estudiados los perezosos siempre formaron
parte de la dieta y fueron la presa más común, los monos podrían ser la presa preferida
en condiciones de alta disponibilidad, como en áreas en donde no sufran presión de
cacería por parte de humanos y su hábitat esté bien conservado.
No se encontraron diferencias en la tasa de aporte de alimento ni en el tipo de presas
entregadas según las diferentes edades de los pollos, a pesar de que en algunos estudios
con aves rapaces se observó que, a medida que las crías se desarrollan y sus necesidades
energéticas aumentan, los padres aumentan su tasa de entrega de alimento o el tamaño
de las presas que capturan. Nuestros datos pueden sugerir que en el área de estudio las
águilas obtienen presas que exceden los requerimientos de los juveniles a lo largo de su
periodo de cría, puesto que el peso y tamaño de las especies capturadas es, en general,
considerablemente alto y suficiente para cubrir sus necesidades.
Todas las especies identificadas que formaron parte de la dieta del águila harpía
están dentro de alguna categoría de amenaza, ya sea a nivel nacional, internacional o
ambos, lo que supone que además de sus propias características biológicas, como la de
criar un único pollo cada tres años, la pérdida de hábitat y la persecución humana, la
estabilidad de las poblaciones de águila harpía se encuentra también amenazada porque
los recursos de los que depende también lo están.
Se calculó que la distancia mínima entre nidos más cercanos es de 4,9 ± 0,7 km,
ocupando cada pareja un área de 19,6 ± 5,7 km2, resultados parecidos a los obtenidos
en Panamá, en donde las condiciones de conservación del hábitat son similares a las de
Ecuador. En general, las especies tropicales poseen requerimientos territoriales menores
que las de las regiones temperadas (Keran 1978), tal y como ocurre con las dimensiones
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del área de cría reportadas para el águila real (Aquila chrysaetos), un águila de gran
tamaño de la región temperada que requiere 285,7 km2 por pareja (Carrete et al. 2001).
Sin embargo, en hábitat alterados como los que ocupan algunas parejas de águila
pitecófaga (Pithecophaga jefferyi) en Filipinas, los tamaños de territorio pueden ser
mayores, puesto que si debido a la alteración del ambiente los recursos que necesita son
más escasos, los territorios se conformarán por áreas más amplias (Bueser et al. 2003).
Aunque teóricamente pueden encontrarse áreas de cría al menos hasta los
1.200 m s. n. m., los nidos encontrados en nuestro estudio se ubicaron a
219  11.7 m s. n. m., ya que la altitud del área de estudio comprende altitudes entre
250 y 326 m s. n. m. A diferencia de Perú, en donde la mayor parte de los nidos que
fueron identificados se hallaron en bosques de Tierra Firme (Giudice 2005), en este
trabajo las áreas de cría se localizaron especialmente en bosques Inundables. Aunque
en algunos estudios se ha encontrado que los bosques de Tierra Firme poseen mayor
riqueza de especies que los anteriores (Balslev et al. 1987, Peres 1997, Patton et al.
2000, Haugaasen & Peres 2005 a, b), parece que la densidad y biomasa total tanto de
primates como de perezosos es menor que en los bosques Inundables (Haugaasen &
Peres 2005 a, b), por lo que éstos ofrecen condiciones favorables para que el águila
pueda obtener suficientes recursos. Por otra parte, los nidos se encontraron en áreas de
baja presión antrópica, aunque éste es un dato esperable debido a que el área de estudio
se sitúa en un área protegida y sus alrededores inmediatos. Asentamientos indígenas se
encontraron muy próximos a las áreas de cría, lo que sugiere que éstos mantienen sus
territorios en condiciones que al menos son aceptables como para que no interfieran con
las necesidades de cría del águila harpía. Así, la presencia de estas comunidades no es
un factor desventajoso para esta especie, siempre y cuando no sufran de persecución
humana.
El hábitat potencial disponible para el águila harpía en Ecuador se estimó en
77.849,86 km2. El rápido avance de monocultivos, de la frontera agrícola y de las
consecuencias sociales y ambientales de la explotación de crudo en la Amazonía
ecuatoriana pone en riesgo la estabilidad de las parejas de águila harpía que se
encuentran en el límite del bosque. Sin embargo, si esto se controla, el futuro de las
poblaciones de esta especie es alentador en esta región del país, pero ocurre lo contrario
al occidente de la cordillera andina, en donde la fragmentación y pérdida de hábitat son
mucho más evidentes.
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Detectar individuos muertos de águila harpía es extremadamente complicado,

especialmente si los ejemplares no cuentan con marcas de identificación o seguimiento,
por lo que es fácil subestimar el número de estos eventos. La mortalidad de adultos
registrada en este estudio (9,4%) fue menor que la mortalidad juvenil (28,6%) y, en
todos los casos, debida a persecución humana. Miedo a la especie, consideración de
depredadora de animales domésticos, uso de sus elementos corporales con motivos
mágicos y venganzas personales fueron las razones detectadas para justificar los
disparos que sufrieron los distintos ejemplares, entre los que se incluye un juvenil. Los
conflictos antropogénicos juegan un papel importante en el declive de algunas
poblaciones de rapaces (Raptor Research Foundation 2016), y está considerada como
una de las causas de disminución de la especie estudiada (Vargas et al. 2006)
La mortalidad juvenil en este estudio es menor a la observada para 13 especies de
aves rapaces europeas (Newton 1979), aunque no se conocen estudios que cuantifiquen
este aspecto en este grupo y en la región Neotropical. El momento de aprendizaje del
vuelo es un periodo crítico para la supervivencia de los juveniles, puesto que durante
éste pueden ocurrir accidentes que provoquen su muerte. Tal y como también se
observó en Venezuela (Álvarez-Cordero 1996), durante esta etapa, adoptan posturas que
les desequilibran y, si no son capaces de corregirlas, pueden caer al estrato más bajo del
bosque con el riesgo de quedarse atrapados en éste y no poder volver a ascender, tal y
como probablemente ocurrió en cuatro de las seis muertes descubiertas.
Debe continuarse el seguimiento de adultos y juveniles para determinar con mayor
certeza las principales causas de su muerte, ya que conocer cuáles son los elementos que
afectan a su éxito reproductor es esencial para entender la dinámica poblacional de esta
especie y, de esta manera, desarrollar estrategias de conservación más eficientes.
Los esfuerzos para conservar no solo al águila harpía, sino a la biodiversidad
existente en el área donde se encuentra, deberían estar integrados con un desarrollo
social y económico respetuoso con el ambiente y con las necesidades locales, haciendo
partícipe a la población en los proyectos que se ejecutan en sus distintas etapas y
ejerciendo control sobre la utilización de recursos naturales a través de incrementar la
voluntad política para ello. La educación ambiental y la sensibilización tanto a nivel
local como nacional son elementos necesarios e ineludibles para disminuir los conflictos
gente-fauna y para obtener el apoyo social que requieren las acciones de conservación.
El concepto de “Especie Bandera” puede ser una forma útil de aumentar el alcance e
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interés de la población acerca de la importancia de conservar la biodiversidad,

pudiéndose obtener resultados favorables para emprender acciones que la beneficien.
1.6. Conclusiones generales de la Tesis Doctoral.
1. Se describen por primera vez los movimientos de juveniles de águila harpía

utilizando telemetría vía satelital con GPS. Esta especie posee un prolongado
periodo de dependencia juvenil que se extiende hasta al menos los dos años de
edad, en donde sólo ocurren desplazamientos cortos dentro del territorio paterno
sin evidenciarse tendencia dispersiva alguna.
2. Se estima en 28 meses la edad en la que un juvenil comienza su dispersión

juvenil. Antes de este momento, las distancias que recorren son cortas y siempre
dentro del área natal.
3. En sus movimientos exploratorios de dispersión, el juvenil recorre una distancia

media de 1,07 km al día, llegando a alejarse como máximo 35,1 km de su área
natal a los 35 meses de edad, mientras que para otras especies de águilas estas
distancias son considerablemente mayores.
4. El águila harpía presenta hábitos tróficos especialistas aunque es capaz de predar

sobre cierta diversidad de especies que habitualmente se encuentran en el estrato
más alto del bosque. Durante el periodo de cría, los machos son los que más
frecuentemente acuden con alimento al nido y ambos padres se presentan
preferentemente por las mañanas y cada 3,8 días de media.
5. Los perezosos son las presas más seleccionadas por los adultos. Sin embargo, en
áreas en donde existe menos presión de cacería de primates, los parentales
aportan en los nidos más monos que perezosos pudiendo ser los primeros la
presa preferida en condiciones de alta disponibilidad de este recurso.
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6. Existen condiciones adecuadas para que el águila harpía establezca su área de

cría desde 0 hasta al menos 1200 m s. n.m. en Ecuador siempre y cuando el
bosque se encuentre en buen estado.
7. La densidad de áreas de cría de águila harpía en nuestra área de estudio es

relativamente alta, estimando una densidad de 5 nidos/100 km2 en esta región
amazónica y, por el momento, de forma similar a la densidad existente en
Panamá, aunque mayor información se requiere para el continente
Suramericano.
8. Los bosques Inundables en buen estado ofrecen condiciones favorables para que
sirvan como áreas de nidificación de águila harpía, siendo el tipo de bosque en
donde más frecuentemente se localizaron los nidos en este estudio.
9. El éxito de cría de águila harpía puede ocurrir cerca de comunidades humanas

que conservan sus territorios y no tienen conflictos con esta especie.
10. El futuro del águila harpía en la región oriental de Ecuador cuenta con buenas
previsiones siempre y cuando se mantengan controladas las presiones de pérdida
de hábitat que comienzan a afectar a esta parte del país. En la región occidental
su situación se encuentra más comprometida debido a la desaparición de la
mayor parte de su hábitat original. En total, estimamos 77.849,86 km2 de hábitat
potencial para esta especie en Ecuador.
11. Se detectó que la etapa juvenil es la de mayor mortalidad, superior a la del

estado adulto y debida sobre todo a causas naturales. El momento de aprendizaje
del vuelo es un periodo crítico para la supervivencia de los juveniles puesto que
durante éste pueden ocurrir accidentes que provoquen su muerte.
12. Los casos registrados de mortalidad adulta se originaron por disparos a causa de
temor hacia la especie, de su consideración como depredadora de animales
domésticos, utilización de partes de su cuerpo como elementos mágicos y
elemento con el que saldar venganza de conflictos intracomunitarios.
33
13. Las áreas de cría deben ser tenidas en cuenta en una estrategia de conservación
no sólo durante el tiempo en el que el pollo se encuentra en el nido, sino al
menos un año después de su nacimiento.
14. La educación ambiental y la sensibilización son acciones fundamentales para

disminuir los casos de mortalidad no natural del águila harpía. Los casos de
mortalidad juvenil debido a accidentes pueden reducirse con un monitoreo más
acentuado durante esta etapa. Utilizar a esta especie como “Especie Bandera”
puede favorecer el acercamiento a la sociedad de las consecuencias a las que se
enfrenta la biodiversidad a causa de los problemas de conservación que existen
en las diferentes áreas del país.
15. Comprender mejor las circunstancias, intereses, conflictos y cultura de las

comunidades que conviven con los recursos naturales y aumentar su
participación en todas las fases de desarrollo de las distintas iniciativas de
conservación puede ser una estrategia útil para aumentar las posibilidades de
alcanzar los objetivos de esas intervenciones.
16. Una mayor cantidad de estudios son necesarios para comprender la biología y
las necesidades de esta especie y para poder concretar de la forma más adecuada
posible los mecanismos apropiados para su conservación.
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43
CAPÍTULO 2
Juvenil de águila harpía en la Reserva Cuyabeno. Foto: R. Muñiz López
Los espíritus que protegen a los Cunsi Pindo:

El águila harpía (Cunsi pindo en lengua Aí o Cofán) suele criar en grandes árboles, uno de ellos el
llamado Ceibo (Ceiba pentandra). Según los Cofanes, en cada Ceibo vive un espíritu, llamado
“Atsatábahe Kukuya”, y cuando ese espíritu abandona su árbol éste comienza a marchitarse hasta morir.
“El Atsatábahe Kukuya protege y mezquina al águila que esté criando sobre esta especie, impidiendo
cualquier daño que los humanos pudieran hacerle. Así, cuando necesiten capturar un pichón de Águila
harpía para colocarle su “mochila” (refiriéndose al transmisor satelital-GPS) necesitan pedir permiso al
espíritu del Ceibo para que éste comprenda y autorice nuestra tarea” (Osvaldo Criollo, indígena Aí /
Cofán, com. pers. 2005)
45
46
CAPÍTULO 2
MOVIMIENTOS DE ÁGUILAS HARPÍAS Harpia harpyja DURANTE LOS DOS

PRIMEROS AÑOS DESDE SU NACIMIENTO
RUTH MUÑIZ-LÓPEZ1, 2, RUBÉN LIMIÑANA1, 3, GONZALO D. CORTÉS4, 5 and

VICENTE URIOS1
1
Estación Biológica Terra Natura, Instituto Universitario de Investigación CIBIO, Universidad
de Alicante, Apdo. correos 99, E-03080 Alicante, Spain;
2
Sociedad para la Investigación y Monitoreo de la Biodiversidad Ecuatoriana (SIMBIOE), C/
Jorge Juan N31-120 y Mariana de Jesús, Quito, Ecuador;
3
Instituto de Investigación en Recursos Cinegéticos (IREC), CSIC-UCLM-JCCM, Ronda de
Toledo, s/n, E-13005 Ciudad Real, Spain;
4
Área de Biodiversidad y Conservación, Museo Nacional de Historia Natural, DICYT/MEC,
Casilla de Correo 399, CP 11000, Montevideo, Uruguay and
5
Laboratorio de Etología, Ecología y Evolución n, Instituto de Investigaciones Biológicas
Clemente Estable, MEC, Montevideo, Uruguay.
Este capítulo reproduce íntegramente el texto del siguiente manuscrito:
Muñiz-López Ruth, Rubén Limiñana, Gonzalo D. Cortés and Vicente Urios. 2012.
Movements of Harpy Eagles Harpia harpyja during their first two years after hatching.
Bird Study 59 (3): 1-6.
Resumen
Dos águilas harpías juveniles fueron equipadas con transmisores satélite cuando se
encontraban en sus nidos. Durante los dos primeros años los desplazamientos de ambas
aves ocurrieron hasta distancias muy cortas, registrándose una distancia máxima de 1.3
km desde el nido. Durante el periodo de estudio, ninguna de las dos comenzó la
dispersión juvenil pero en una de ellas se detectó un incremento en el distanciamiento
medio mensual al nido con la edad, mientras que para el otro individuo esto no ocurrió.
Proteger los territorios de los adultos de águila harpía puede favorecer así mismo la
protección de los juveniles durante un largo periodo e tiempo.
47
MOVEMENTS OF HARPY EAGLES Harpia harpyja DURING THEIR FIRST

TWO YEARS AFTER HATCHING
RUTH MUÑIZ-LÓPEZ1, 2, RUBÉN LIMIÑANA1, 3, GONZALO D. CORTÉS4, 5 and

VICENTE URIOS1
1
Estación Biológica Terra Natura, Instituto Universitario de Investigación CIBIO, Universidad
de Alicante, Apdo. correos 99, E-03080 Alicante, Spain;
2
Sociedad para la Investigación y Monitoreo de la Biodiversidad Ecuatoriana (SIMBIOE), C/
Jorge Juan N31-120 y Mariana de Jesús, Quito, Ecuador;
3
Instituto de Investigación en Recursos Cinegéticos (IREC), CSIC-UCLM-JCCM, Ronda de
Toledo, s/n, E-13005 Ciudad Real, Spain;
4
Área de Biodiversidad y Conservación, Museo Nacional de Historia Natural, DICYT/MEC,
Casilla de Correo 399, CP 11000, Montevideo, Uruguay and
5
Laboratorio de Etología, Ecología y Evolución n, Instituto de Investigaciones Biológicas
Clemente Estable, MEC, Montevideo, Uruguay.
This chapter reproduced entirely the text of the following manuscript:
Muñiz-López Ruth, Rubén Limiñana, Gonzalo D. Cortés and Vicente Urios. 2012.
Movements of Harpy Eagles Harpia harpyja during their first two years after hatching.
Bird Study 59 (3): 1-6.
Capsule: Two juvenile Harpy Eagles were tagged at their nests with satellite
transmitters. During the first two years after hatching both birds moved very short
distances, with a maximum recorded distance of 1.3 km from the nest. During the study
period, neither bird started juvenile dispersal, but while one of the birds showed a
significant overall increase in mean monthly distance from nest with age, the other bird
did not. Protecting territories of adult Harpy Eagles may enhance the protection of
juveniles for a long period of time.
In large, long-lived raptors with delayed (Ferrer 2001, Whitfield et al. 2004,
maturity, the period between fledgling Penteriani et al. 2006, 2011, Soutullo et
and recruitment to the breeding al. 2008a). However, because of the
population may take several years and is difficulty of obtaining adequate data
usually poorly understood (Whitfield et during these periods, little is known on
al. 2009). Juvenile dispersal represents the juvenile stage of most raptor species
the movements undertaken by juveniles, (Penteriani & Delgado 2009).
once they become independent from The Harpy Eagle Harpia harpyja is
their parents, to find a breeding site one of the largest raptors of the
(Clobert et al. 2001). American continent, being distributed
The onset of juvenile dispersal is from southern Mexico to northern
not easy to detect and usually it has Argentina (Ferguson- Lees & Christie
been more or less arbitrarily defined. 2001). Currently, the species is
Nevertheless, a simple and adequate threatened across its whole distribution
approach to estimate the onset of range and population numbers are
dispersal considers that this stage of declining, mainly due to human
bird’s life-cycle starts when juveniles persecution and habitat loss and
abandon their parental territory fragmentation (Vargas et al. 2006,
(sometimes based on when juveniles BirdLife International 2012). For this
reach the mid-distance between adjacent reason, the Harpy Eagle is globally
nests in the region and remain beyond listed as Near Threatened (BirdLife
that distance for a given period of time; International 2012). In Ecuador (where
e.g. Ferrer 1993, Soutullo et al. 2006a, this study was conducted) it is listed as
Cadahía et al. 2008). The period Vulnerable (Granizo et al. 2002).
between the first flights and the onset of Although several aspects of its
juvenile dispersal is known as the post- biology and ecology have been studied
fledgling dependency period, when at several areas across its distribution
juveniles are still dependent on their range, such as Brazil (e.g. Galetti & de
parents (e.g. Soutullo et al. 2006a). The Carvalho Jr 2000, Lenz & Marajo dos
fate of juvenile birds during these Reis 2011), Ecuador (Muñiz-López
periods may have important 2007, 2008), Panama (e.g. Vargas &
consequences for population trends and, Vargas 2011), Peru (Piana 2007) and
thus, they are of particular importance Venezuela (Álvarez-Cordero 1996), the
from a conservation point of view Harpy Eagle is one of the least-known
raptor species of the world. Most of the trees, and being fully dependent on
information available on the species is parents in terms of feeding (Álvarez-
related to diet and adult behaviour at Cordero 1996). Harpy Eagles are
their breeding areas (Rettig 1978, supposed to be in this post-fledgling
Álvarez-Cordero 1996, Muñiz-López dependency period for a long time
2008) and recently also on population (Álvarez-Cordero 1996), although no
genetics (Banhos et al. 2008, Lerner et quantitative data to confirm this have
al. 2009). The species feeds primarily been obtained to date.
on tree-dwelling mammals, particularly In recent years, the use of satellite
monkeys and sloths, as well as on large telemetry has become widespread and
birds and reptiles (Fowler & Cope 1964, predominant in the study of migratory
Rettig 1978, Álvarez-Cordero 1996, movements of several raptor species
Galetti & de Carvalho Jr 2000, Piana (e.g. Limiñana et al. 2007, 2012a,b,
2007, Muñiz-López 2008, Springer et Strandberg et al. 2010, López-López et
al. 2011). al. 2010, Mellone et al. 2011), as well as
The Harpy Eagle has probably the in detailed studies of juvenile dispersal
longest breeding period of any raptor, and habitat use (e.g. Urios et al. 2007,
usually breeding once every two and a Soutullo et al. 2008b, Cadahía et al.
half or three years (Rettig 2010). Here, we use GPS-satellite
1978,Álvarez-Cordero 1996). telemetry to describe the movements of
According to the few studies conducted two freeranging juvenile female Harpy
on this species, adult home-ranges are Eagles in Ecuador during the first two
generally large, ranging between 16 and years after hatching.
79 km2 (Álvarez-Cordero 1996, Muñiz- This study was conducted within
López 2007, Vargas & Vargas 2011). the reserve ‘Reserva de Producción
Juvenile Harpy Eagles have Faunística Cuyabeno (-0.117° N, –
rarely been monitored and the only data 75.833°E, northeastern Ecuador). This
regarding their first movements suggest reserve is dominated by rainforest,
that fledgling occurs when birds are although habitats range from evergreen
between 120 and 160 days old (Rettig forests to flooded lowlands (Cerón et al.
1978, Álvarez-Cordero 1996, Muñiz- 1999, Palacios et al. 1999). There, two
López 2007). After that, juveniles are juvenile Harpy Eagles (‘Masakay’ and
mostly inactive, making only short ‘Tava’) were trapped at their nests.
flights within the nest-tree or nearby ‘Masakay’ was tagged on 15 July 2006
50
in its third month after hatching, and During the study period, a total of
‘Tava’ on 28 March 2009 in its seventh 7170 GPS locations were obtained for
month after hatching. Age of birds at ‘Masakay’ and 6337 for ‘Tava’.
tagging was known because the nests Locations were transformed from
were monitored before the eggs hatched Geographic Coordinate System to UTM
by Cofanes natives, who collaborated in coordinates for calculations. For each
the project and recorded hatching date. individual, we calculated the
Both birds were sexed as females loxodromic distance from every
by means of body size and weight recorded position to its nest. We
measures, according to reference values evaluated the relationship between the
for juveniles in del Hoyo et al. (1994). distance to nest and age of birds
Eagles were tagged with 70-g (months after hatching) using linear and
Argos/GPS satellite transmitters quadratic regression analysis (distance
(Microwave Telemetry Inc.), which to the nest ~ bird age × bird identity +
were affixed to their backs using a bird age squared × bird identity). Since
Teflon harness. The full transmitter these initial models presented
equipment did not exceed 1.5% of the heterogeneity of variances, we then
juveniles’ body mass, which is conducted Generalized Least Square
considerably below the 3% suggested to (GLS) regression models to overcome
minimize the effects of additional mass this analytical issue (Pinheiro & Bates
on birds’ movements (Kenward 2001). 2000, Zuur et al. 2010). Therefore,
Satellite transmitters were programmed considering the possible existence of
to record one GPS position (nominal proportional or potential structures of
accuracy + 18 m) every hour between variances and also considering that
09:00 and 03:00 for ‘Masakay’ and different variance structures could
between 11:00 and 03:00 for ‘Tava’ emerge for each bird, these alternative
(local time). Data were recorded until variance structures were evaluated for
both individuals reached the age of two each one of the independent variables
years (i.e. the last day of their 23rd (Zuur et al. 2009). In order to find the
month after hatching, taking into optimal variance structure we computed
account age of birds at tagging). For the same model using different variance
both birds, we excluded data from the structures and compared each GLS
tagging month from the analyses, model using Akaike’s Information
because it was only partially recorded. Criterion (AIC) (Burnham & Anderson
51
2002, Zuur et al. 2009). All these

models were fitted with the gls function
from the NLME package (Pinheiro et al.
2011) for R software (R Development
Core Team 2011). When doing this, the
best model (lowest AIC value) was the
one that included the second-order
polynomic variable (distance to nest ~
bird age × bird identity + bird age

squared × bird identity) and included a
power variance structure (varPower ~
Figure 1. Movements of two Harpy
bird age|bird identity, in R), thus taking Eagles (A: ‘Masakay’ and B: ‘Tava’),
into account differences in monthly tracked by satellite telemetry during
spread of data for each bird. As two years after hatching. Open dots
interactions in this model were represent every recorded distance to
significant (P < 0.001 in all cases), these nest forevery month within this two-
results show that during the study year period. The line represents the
period the two tracked juvenile eagles quadratic model that better described
exhibited significantly different the movements of both eagles in the
behaviours. study period (see text for details and
parameter estimates).
To deconstruct the movements

pattern of each individual, we
conducted GLS models separately to
test if distance to nest showed a linear
or quadratic trend for each individual
(distance to nest ~ bird age + bird age
squared), also accounting for the
observed heterogeneity of variances.
The best individual GLS models
(according to AIC) included the age
squared term, but a different variance
structure. The selected model for
52
‘Masakay’ included a power variance ranges and breeding territories, this

structure of the quadratic covariate process may take several years until a
(varPower ~ bird age + squared bird age, breeding territory is occupied (e.g.
in R) (parameter estimates: intercept = Urios et al. 2007, Cadahía et al. 2009,
16.1, P <0.001; age = 2.7, P < 0.001; Whitfield et al. 2009). Given the extent
age squared = –0.1, P < 0.001). The of the adult home-ranges in the Harpy
selected model for ‘Tava’ included a Eagle, ranging between 14 and 79 km2
variance structure proportional to the (Álvarez-Cordero 1996, Vargas &
quadratic covariate (varFixed ~ bird age Vargas 2011), and considering the
+ squared bird age, in R) (parameter movements reported here, it is clear that
estimates: intercept = 110.0, P < 0.001; none of the tracked juveniles left the
age = –10.6, P < 0.001; age squared= parental territory during the study
0.8, P< 0.001). These differences in period. Therefore, the onset of the
parameter estimates revealed different juvenile dispersal in the species does
individual trends in distance to nest not take place within the first two years
during the study period (Fig. 1). after hatching, and the stage of life
Whereas ‘Tava’ showed an overall reported here corresponds to the post-
increasing trend in its monthly distances fledgling dependence period (Soutullo
to nest, ‘Masakay’ first showed a slight et al. 2006a, Cadahía et al. 2008).
increase in distances from the nest but Similar results for the Harpy Eagle
then it began to reduce the extent of its were found by Álvarez- Cordero (1996),
movements from the nest, spending who reported distances to the nest of
more time closer to the nest (Fig. 1). 300–600 m for juveniles 2 years old;
During the study period, maximum however, the data of Álvarez-Cordero
distance to the nest achieved by ‘Tava’ (1996) are based on direct observations,
was 1300 m when it was 23 months old, which, in the closed-canopy forests
whereas for ‘Masakay’ this maximum where Harpy Eagles live, may have
distance was 296 m at the age of 13 precluded getting observations at larger
months (Fig. 1). distances from nest.
Juvenile dispersal is the ecological The individual eagles showed a
process that leads juvenile birds to find different behaviour during the study
a site for breeding (Clobert et al. 2001). period. ‘Tava’ showed a progressive
In large bird species with delayed distancing from nest during the study
maturity and relatively large home- period, while ‘Masakay’ first showed a
53
distancing from nest but then began to survival probability (Skutch 1976) or to
spend more time near the nest (see Fig. be in better condition at the onset of
1). dispersal (Young 1996).
Progressive distancing from the nest Age of first reproduction for Harpy
during the post-fledgling dependency Eagles breeding in captivity has been
period is common in several large estimated as 4 years; however, taking
raptors (e.g. O’Toole et al. 1999, into account that the onset of juvenile
Soutullo et al. 2006a,b, Cadahía et al. dispersal for freeranging individuals in
2008). However, during this undisturbed environments takes place at
dependency period, juvenile birds are ages older than 2 years, age of first
mostly fed by parents, which may reproduction in the wild is likely to take
explain the constant return of both birds place at ages probably much older than
to their nests observed here. However, 4 years (see also Shaner 2011), which
during this period, juvenile raptors also would represent one of the longest
make their first hunting attempts within dispersal periods among raptors.
parental territories (e.g. Ferrer 1993, However, as Harpy Eagles live in
Kitowski 2009), and this has been also habitats that are largely homogeneous
observed for Harpy Eagles (Muñiz- and undisturbed, there may be little
López 2007). Therefore, differences in need to cover very large distances to
prey abundance or availability in the find a breeding site, contrary to other
areas used by these Harpy Eagles may species in temperate heterogeneous
explain, at least partially, the different habitats (e.g. Urios et al. 2007, Cadahía
behaviour of these individuals, with the et al. 2009, Whitfield et al. 2009),
eagle moving closer to its nest possibly which would reduce the overall time
being in a less-productive area. spent dispersing.
Large, tropical birds usually show Future tracking studies of
reduced clutch sizes and long breeding individuals should focus on these
seasons due to the relative aspects, because this floating period of
environmental stability of their habitats large raptors is of great importance from
(Russell & Rowley 2000). As a a conservation point of view (Penteriani
consequence of the reduced brood size, et al. 2006, 2011, Soutullo et al. 2008a).
parental care is usually longer and Our results and conclusions are
juveniles spend more time at the limited because of the small sample size
parental territory to enhance their that is often associated with such
54
satellite-tracking studies. However, this Acknowledgements

approach has the advantage of obtaining
spatially explicit information about We are indebted to Fundación Terra
raptors’ movements, which is of great Natura for funding the Project on
importance from a conservation point of satellite tracking of Harpy Eagles in
view, even if only a few birds are Ecuador. We would like to thank
tracked (e.g. Urios et al. 2010). In the Alexander Blanco for his support and
case of the Harpy Eagle, this is help in capturing wild Harpy Eagles.
exacerbated by the secretive nature of We are also thankful to the indigenous
the species, the difficulties in finding communities of the study area for their
their nests, as well as carrying out the essential work with the species. The
fieldwork in remote dense forests. Environment Ministry of Ecuador and
Results presented here indicate that EcoFondo funded the fieldwork. A.
territories occupied by adults are also Soutullo and P. López-López provided
important for juvenile birds for at least insightful comments on an earlier draft
2 years after hatching. Therefore, of the manuscript. We are also grateful
conservation actions at these areas to P. Whitfield and W. Cresswell for
should not only be focused during the their comments, which substantially
pre-fledgling period, but also improved the manuscript. R. Limiñana
throughout the year, given that the had a postdoctoral grant (reference
juveniles are actively using the same 10/12-C) co-funded by ‘Consejería de
areas used by parents for a long time. Educación y Ciencia’ (JCCM) and the
Juvenile survival is one of the most European Social Fund. Gonzalo D.
important parameters determining Corte´s is supported by a ‘Beca de
population trends in long-lived raptors Maestría’ of the Sistema Nacional de
(Soutullo et al. 2008a). Hence, Becas of the Agencia Nacional de
protecting these adult territories would Investigación e Innovación (reference
also not only directly affect adult POSNAC 2011 POS-2011-1-3383).
survival but also juvenile survival, This article is part of the PhD thesis of
which would be of paramount R. Muñiz-López at the University of
importance for conservation of the Alicante.
Harpy Eagle.
55
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60
CAPÍTULO 3
“Tava”, hembra juvenil de Águila harpía que fue equipada con emisor satélite. Foto: A. Blanco /PCAHE-
SIMBIOE
Águila que va volando, no dice a dónde… ni cuándo.
Anónimo
61
62
CAPÍTULO 3
DISPERSIÓN JUVENIL DEL ÁGUILA HARPÍA Harpia harpyja
VICENTE URIOS, RUTH MUÑIZ-LÓPEZ & JAVIER VIDAL-MATEO
Palabras clave: rapaces, telemetría satélite, ecología del movimiento, agresión parental.
Resumen
Los movimientos de dos águilas harpía juveniles antes y durante el periodo de
dispersión son estudiados mediante telemetría satelital GPS. Una de las harpías
comienza la dispersión durante el mes 28 de edad, mientras que la otra en esas fechas
permanece en las cercanías del nido y fallece a los 30 meses de edad. El juvenil que se
dispersa se aleja un máximo de 35.1 km durante el periodo de estudio y ocupa un área
de 386 km2 según kernel fijo del 95%.
63
JUVENILE DISPERSAL OF HARPY EAGLE Harpia harpyja
VICENTE URIOS, RUTH MUÑIZ-LÓPEZ & JAVIER VIDAL-MATEO
Key words: raptors, satellite telemetry, movement ecology, parental aggression.
Capsule: Movements of two juvenile Harpy Eagles before and during the dispersal
period were studied by GPS satellite telemetry. One of the eagles started the dispersal
during the month 28 of life, while on the same dates the other juvenile remained in the
surrounding area of the nest, suffering parental attacks that might cause its death.
64
The Harpy Eagle Harpia harpyja is one movement ecology is still poorly known
of the largest and heaviest birds of prey (Muñiz-López et al. 2012), in particular
in the world (Brown & Amadon 1968). its juvenile dispersal period. Juvenile
This species usually lives in lowland dispersal comprises the movements
rainforests, with a distribution range undertaken by juveniles in their search
from southern Mexico to northern for a breeding area once they are
Argentina discontinuously (Ferguson- independent from their parents
Lees & Christie 2001). (Greenwood & Harvey 1982, Clobert et
Due to habitat loss and human al. 2001). In large raptors this process
persecution, the populations of this may take several years (Cadahía et al.
eagle are declining (Alvarez-Cordero 2009, Whitfield et al. 2009), and it is
1996, Vargas et al. 2006, BirdLife preceded by a period of parental
International 2016). Its long breeding dependence called post-fledging period.
interval, not studied in depth and This post-fledging period takes place
estimated between two and a half and from the first flights of the bird until the
three years (Rettig 1978, Álvarez- onset of the dispersal (Soutullo et al.
Cordero 1996), makes it even more 2006). The onset of dispersal period is
vulnerable. Currently, for these reasons, not easy to detect, and it has been seen
it is considered a Near Threatened that it can be influenced by a reduction
species by the International Union for of the parental investment in their
Conservation of Nature and Natural offspring (e.g. decreasing the food
Resources (IUCN 2013), and in supply), even with presence of parent-
Ecuador it is included within the offspring aggressions (Alonso et al.
category Vulnerable (Granizo et al. 1987) or between siblings (Holleback
1997). 1974).
Different aspects of the biology of Here we study the movements
the Harpy Eagle have been studied, as before and during the dispersal period
its trophic ecology (Piana 2007, Lenz & of two juveniles Harpy Eagles tracked
Marajó dos Reis 2011, Aguiar-Silva et by GPS satellite telemetry.
al. 2014) or its breeding behaviour Two juvenile female Harpy Eagles
(Rettig 1978, Álvarez-Cordero 1996, were trapped and tagged: ‘Masakay’ (#
Muñiz-López 2007, Rotenberg et al. 49178) and ‘Tava’ (# 48179). Both
2012). However, due to the difficulty of eagles were tagged at their nest tree (for
observation of this species, its more details of the capture see Muñiz-
65
López et al. 2012), in the ‘Reserva de Harpy Eagles until they reached the age
Producción Faunística Cuyabeno’ of two years. In this study we track their
(Sucumbíos province, northeastern movements since that date (first day of
Ecuador). Located in the Amazon the 24th month of age) onwards: in
region of Ecuador, this is one of the ‘Masakay’ until month 27 and month 39
most biodiverse places on the planet, in ‘Tava’ (4 and 15 months of tracking
dominated by primary lowland tropical respectively).
moist forest between 250 and 300 m, During the months of study, a total
and flooded tropical evergreen forests of 1320 locations for ‘Masakay’ and
(Cañadas 1983, Stotz et al. 1996, Cerón 3761 for ‘Tava’ were received. Data
et al. 1999, Palacios et al. 1999). were retrieved and managed by Satellite
Eagles were tagged with 70 g Tracking and Analysis Tools (STAT;
Argos/GPS satellite transmitters Coyne & Godley 2005). The distances
(Microwave Telemetry Inc.), which covered every day for each eagle were
were affixed to their backs using a calculated, selecting a location per day
Teflon harness. This equipment did not and trying to approach as much as
exceed 1.5% of body mass, which is possible to midnight. We also calculate
within the recommended limits the distance to the nest from every
(Kenward 2001). One GPS position recorded position.
(nominal accuracy ± 18 m) was Using the Mann-Whitney U test, we
recorded every hour between 09:00 and evaluated the differences in these daily
03:00 (local time) for ‘Masakay’ and distances and distances to nest between
between 11:00 and 03:00 for ‘Tava’. ‘Masakay’ and ‘Tava’ during the
‘Masakay’ was tagged on 15 July 2006 months that data are available for both
in its third month of age, and ‘Tava’ on birds. In the case of ‘Tava’ we also
28 March 2009 in its seventh month of analyzed the differences between the
age. In addition, in collaboration with daily distances before the onset of
the natives of nationality A'i / Cofan, an dispersal when he was in the natal area,
observation tower 30 m high was built, and the distances covered once started
which allowed also take visual records the dispersal period. Daily distances
of the eagles. calculated between non-consecutive
Muñiz-López et al. (2012) days (for lack of data) were excluded
described the movements of these two from the analysis.
66
In addition, with all obtained GPS coordinates to an Equal-Area

locations, we estimated the hourly Cylindrical projection using the
distances as the straight distance Projector! Extension for ArcView 3.2.
between two locations corresponding to We also calculated the Minimum
consecutive hours. These distances Convex Polygon (MCP) encompassing
could be calculated between 09:00 and all the locations obtained for the
3:00 hours (local time), the time period different periods (post-fledging and
in which the locations were obtained. dispersal period), performing the same
We used the Kruskal-Wallis test to transformation as for the kernel
examine whether there were differences analyses.
in the hourly distances according to the The individual eagles showed a
time throughout the day, and the different behaviour. ‘Tava’ remained in
multiple comparison Games-Howell test the natal area until the beginning of the
to check whether there was any peak of month 28 of age: on 6 December 2010
activity. began to move away from the nest
All statistical analyses were towards the North and started its
performed with IBM SPSS Statistics juvenile dispersal (Fig. 1). All available
ver. 22.0. Significance level was GPS data of ‘Masakay’ (from month 24
established at P < 0.05. of age to 27) were located in the natal
We estimated the home-range of area, without leaving it. Coinciding with
both Harpy Eagles during the period in the second month of dispersal of ‘Tava’,
the natal area and during the dispersal ‘Masakay’ in the middle of the month
period using a fixed kernel approach 29 of age begins to suffer attacks by one
(Worton 1989), including all GPS of the parents, presumably the male, as
locations. We calculated the 95%, 75% we became aware thanks to the visual
y 50% fixed kernels using the Animal observations carried out from the tower.
Movement extension for ArcView 3.2 These attacks consisted in quick flights
(Hooge & Eichenlaub 1997). showing the claws and approaching the
We used the least squares cross juvenile who was perched on the branch
validation (LSCV) procedure to of a tree adjacent to the nest tree.
calculate the smoothing parameter H ‘Masakay’ was found dead on the next
(Silverman 1986). To estimate the real month (October 2008) a few meters
size of home-ranges we transform the from the nest tree. Furthermore, the
kernel polygons in geographic daily distances covered by ‘Tava’ and
67
‘Masakay’ during the months for which The hourly distances changed
data are available for both birds (in their significantly throughout the day (χ2 =
natal area) were significantly different 65.61, df = 11, P < 0.000), although
(Z = –10.72, P < 0.000), making shorter there was no a prominent peak of
displacements ‘Masakay’: 0.03 km on activity, moving on average 0.11 (±
average (± sd) compared to 0.48 km of 0.14) km/h. The maximum distance
‘Tava’. In the same way it happens with recorded in one hour was 0.99 km,
the distances to the nest measured for although in 95% of the hourly
each day (Z = –34.29, P < 0.000): movements it was below 0.42 km.
‘Masakay’ moved away an average There was activity during all daylight
distance of 0.03 km and ‘Tava’ 0.35 km hours.
before the onset of the dispersal (their During the time that ‘Tava’
maximum distance were 0.24 and remained in the natal area (zone A in
2.2 km respectively; Fig. 2). Fig. 1), it occupied an area of 0.387 km2
In the case of ‘Tava’, the daily according to 95% fixed kernel (0.071
distances covered before and after the and 0.031 km2 for 75% and 50% kernels
onset of the dispersal were significantly respectively, and 6.064 km2 for MCP).
different (Z = –8.72, P < 0.000), with an The estimated area during the eleven
average of 0.48 (± 0.46) km while the months of dispersal (zone B in Fig. 1)
eagle remained in the natal area and that have been studied was 386 km2
1.47 (±0.90) km during the dispersal. according to 95% fixed kernel (137 and
The maximum daily distance recorded 42 km2 for 75% y 50% kernels, and 579
was close to 4 km, although during the km2 for MCP). The 95% kernel for
dispersal period it was less than 2 km in ‘Masakay’ encompassed 2783 m2 (793
most cases (69.03%). Only in 3.87% of and 370 m2 for 75% and 50% kernels,
the daily segments, the distance and 0.112 km2 for MCP).
exceeded 3 km. During the study period The Harpy Eagle presents the
‘Tava’ moves away from the nest a highest record of permanence in the
maximum of 35'1 km (in the 35th natal area and duration of post-fledging
month of age; Fig. 2). dependence period between raptors
The hourly distances were only together with the Philippine Eagle
calculated for ‘Tava’ because Pithecophaga jefferyi (del Hoyo et al.
‘Masakay’ remained static most of the 1994, Álvarez-Cordero 1996, Muñiz-
time making only short displacements. López et al. 2012). According to our
68
data, although we only based on is not common in all raptors

tracking of two individuals, the Harpy (Bustamante & Hiraldo 1990,
Eagle begins its juvenile dispersal Bustamante 1994, Boileau &
period and left its natal area around the Bretagnolle 2014), parental aggression
28th month of life. toward the juveniles has been observed
This age is greater than that has in other species (Alonso et al. 1987).
been seen in other eagles in which was This conflict between adults and
studied the end of the post-fledging offspring (Trivers 1974), promoted by
dependence period (Wood et al. 1998, parents that reduce the food supply or
Soutullo et al. 2006a, Cadahía et al. increase their aggressive behaviour
2008). There is a coincidence between toward juveniles, favours the
the onset of the dispersal of ‘Tava’ and independence and the onset of the
the aggressive behaviour by one of the dispersal (Hiraldo et al. 1989, Arroyo et
adults that the other juvenile suffers. On al. 2002, Balbontín & Ferrer 2005).
the same dates as ‘Tava’ began to Once initiated the dispersal, ‘Tava’
disperse, ‘Masakay’ remained in the remained north of the natal area and
surrounding area of the nest making exploring the territory without stopping
shorter displacements (Fig. 2). permanently in any zone. At 39 months
old, the transmitter stops emitting
Figure 2: signal, but it is likely to continue these
movements until establishing in a
breeding territory and incorporating into
the breeding population, a process that
could take several years (Urios et al.
2007, Cadahía et al. 2009, Whitfield et
al. 2009).
Although the studies on juvenile
dispersal for large forest eagles are
scarce, estimated dispersal area and the
distances covered in the first months
after the start of the dispersal are lower
In our view, for this reason, than in other species. For instance,
possibly it suffered the parental attacks Golden Eagle Aquila chrysaetos (Haller
that might cause the death. Although it 1994, Soutullo et al. 2006b), Spanish
69
Imperial Eagle Aquila adalberti Although in the future it will be

(González et al. 1989, Muriel et al. necessary to conduct more studies of the
2015), Bonelli’s Eagle Aquila fasciata dispersal of other juvenile Harpy Eagles
(Cadahía et al. 2005, 2010) or Crowned and improve the knowledge of their
Eagle Harpyaliateus coronatus (Urios movements, important implications for
et al. 2014) reach hundreds of km away the conservation of this species can be
from the nest and explore large areas discerned from our results. The reduced
before occupying a territory. clutch size and long breeding period
These differences could be due to (Rettig 1978), together with the long
the homogeneity of the habitat that post-fledging and dispersal period
occupies the Harpy Eagle (Muñiz- underline the importance of directing
López 2008) and allows it to have a conservation plans not only to adults but
suitable conditions and sufficient also to juveniles. Therefore,
availability of preys in a smaller surface understanding the movements of
that in the more heterogeneous habitats juvenile dispersal, a period with
in which these eagles live. Abaño et al. relevant demographic consequences,
(2015) also reported larger movements can be of great importance for the
for the Philippine Eagle Pithecophaga conservation of the species (Clobert et
jefferyi, a species similar to Harpy al. 2001, Soutullo et al. 2008, Penteriani
Eagle in size and biology (del Hoyo et et al. 2011).
al. 1994). However, the results are not The long period spent in the nesting
comparable because they involved area and the dispersal movement
released individuals. The New Guinea located not far away from it, facilitate
Harpy Eagle Harpyopsis novaeguineae, the delimitation of areas for protection
smaller in size, inhabits tropical forests of the Harpy Eagle.
of great homogeneity where it has been
seen that also occupies relatively small Acknowledgements
2
areas of territory, of several tens of km
(Watson & Asoyama 2001). Special thanks are due to Terra
There are no quantitative data for Natura Foundation for funding the
other large forest eagles, as the Crested Project on tracking of Harpy Eagles in
Eagle Morphnus guianensis and the Ecuador. SIMBIOE, the Environment
African Crowned Eagle Stephanoaetus Ministry of Ecuador and EcoFondo
coronatus. funded the fieldwork. We are also
70
thankful to Alexander Blanco and the Arroyo, B.E., De Cornulier, T. &

team of participants and volunteers for Bretagnolle, V. 2002. Parental
their help in the captures, and to the investment and parent-offspring
indigenous communities for their conflicts during the post-fledging period
constant support and for allowing us to in Montagu’s Harriers. Anim. Behav.
work on their territories. J. Vidal-Mateo 63:235-244.
is supported by FPU grant of the
Spanish Ministry of Education Álvarez-Cordero, E. 1996. Biology and
(reference FPU014/04671). This article conservation of the Harpy Eagle in
is part of the PhD thesis of R. Muñiz- Venezuela and Panama. PhD Thesis,
López at the University of Alicante. University of Florida.
Balbontín, J. & Ferrer, M. 2005. Factors

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Figure 1:
76
CAPÍTULO 4
Juvenil de águila harpía alimentándose en la plataforma del nido.

Foto: P. Oxford/PCAHE-SIMBIOE
“Uno de nuestros principales símbolos venerados es ‘kenguiwe’. Ella es admirada por nosotros
tanto por ser majestuosa y avisarnos cuando hay peligro, como por tener extraordinaria
habilidad para cazar a través de su fuerza, velocidad y astucia”.
Manuela Omari Ima Omene, indígena Huaorani de Ecuador*.
* Ima Omene M.O. 2012. Saberes Waorani y Parque Nacional yasuní: plantas, salud y bienestar en la Amazonía del Ecuador.
Iniciativa Yasuní ITT. Ministerio Coordinador de Patrimonio, Ministerio del Ambiente, Programa de las Naciones Unidas para el
Desarrollo (PNUD) y Fondo para el Medio Ambiente Mundial (FMAM). Quito. Ecuador. 118 pp.
Capítulo 4
ECOLOGÍA TRÓFICA DEL ÁGUILA HARPÍA (Harpia harpyja) EN

ECUADOR.
RUTH MUÑIZ-LÓPEZ
Resumen
Se analizó el aporte de presas en siete nidos de águila harpía (Harpia harpyja) en el
noreste y el noroeste de Ecuador entre 2002 y 2010. Se encontraron al menos 18 especies
diferentes pero índices de diversidad relativamente bajos en cada muestra. El índice de nicho
trófico fue relativamente bajo (Bsta= 0.28) lo que denota especialización en la dieta del
águila. Los machos entregaron presas más frecuentemente que las hembras y ambos
prefirieron cazar especies ubicadas en el estrato más alto del bosque. El perezoso de dos uñas
(Choloepus spp.) fue la especie que más contribuyó a la dieta del águila harpía estando
presente en todas las muestras. El peso medio de las especies aportadas fue de 4.01 kg. El
grupo taxonómico de las presas aportadas y la frecuencia de entregas no fueron diferentes a
lo largo de las distintas edades de los pollos, ocurriendo preferentemente entre las 7:00 y las
10:00 am. Todas las especies que forman el espectro de la dieta del águila se encuentran en
alguna categoría de amenaza.
Palabras clave: Águila harpía, dieta, Ecuador, Harpia harpyja, nido.
79
TROPHIC ECOLOGY OF THE HARPY EAGLE (Harpia harpyja) IN

ECUADOR
RUTH MUÑIZ-LÓPEZ
Capsule: We examined prey delivered to seven Harpy eagle (Harpia harpyja) nests in
northeast and northwest Ecuador between 2002 and 2010. At least 18 different species were
found but a relatively low diversity indexes were detected in every sample. For our study area
we found a relatively low niche breath (Bsta= 0.28) that means diet specialization. Males
delivered more prey to the nests than females and both preferred species that were located at
the highest forest stratum. Two-toed sloth (Choloepus spp.) was the species that contributed
most to the Harpy eagle diet and it was always present in all the samples. The mean adult
weight of the prey species delivered was 4.01 kg. The taxonomic group of prey provided and
the frecuency of deliveries were similar for all juvenile age classes and prey delivery occurred
preferably between 07:00 to 10:00 H. We determined that all the species delivered were found
under some degree of threat or conservation pressure at a national level and all but one at an
international level.
Key words: Diet, Ecuador, Harpia harpyja, Harpy eagle, nest.

Introduction (Muñiz-López 2003). Since then, fifteen

more nests have been found in the
The Harpy eagle (Harpia harpyja) Ecuadorian Amazon Basin, but only six
is considered one of the most powerful of them have been monitored to collect
birds of prey in the world, as well as data for this study. One additional nest
one of the largest (Brown and Amadon was discovered at the west of the
1968, Collar 1989). It formerly country and this was the only one found
inhabited tropical and subtropical in that area during this study.
rainforest to about 900 m (Stotz et al. In some of its distribution area there
1996) from southern Mexico to northern are few published records about diet of
Argentina (Hilty and Brown 1986). In this species and most of them are based
Ecuador, the species distribution is upon sporadic observations of Harpy
restricted to several small patches of eagle hunting attempts in the field or
forest in the northwest of the country descriptions of provisioning behaviour
and more consistently with an elevation to nests: British Guiana (Fowler and
range below 400 m a.s.l. in the east, Cope 1964, Rettig 1978, Izor 1985),
where Ecuadorian Amazon Basin starts Peru (Eason 1989, Sherman 1991, Piana
(Guerrero 1997, Ridgely and Greenfield 2007), Brazil (Peres 1990, Galetti et al.
2001, Muñiz-López 2002). The Harpy 1997, Ferrari and Port-Carvalho 2003),
eagle is currently considered Near- Venezuela (Álvarez-Cordero 1996),
threatened throughout its range (Collar Panama (Álvarez-Cordero 1996,
et al. 1994, Birdlife International 2013). Touchton et al. 2002, Aparicio 2003)
In Ecuador it is believed to be a and Belize (Rotenberg et al. 2012).
Vulnerable species (Granizo et al. There is a more comprehensive study
1997). that analyzes the Harpy eagle food
Harpy eagles breed every 2.5 to 3 habits in Brazil (Aguiar-Silva et al.
yr and the resulting offspring, usually 2014). One study examined captive-
one per pair, remains in the natal area bred and released Harpy eagles diet and
for at least two years before dispersing hunting (Touchton et al. 2002; Muela
(Rettig 1978, Ruschi 1979, Álvarez- and Curti 2005). All these studies show
Cordero 1996). The first active nest of a that this eagle feed mainly on arboreal
Harpy eagle to be monitored in Ecuador mammals such as sloths, monkeys,
was found in 2002 in the northeast rodents, carnivores and less frequently
81
on reptiles and birds (Fowler and Cope et al. 1999). The temperature averages
1964, Rettig 1978, Eason 1989, 25ºC and environmental moisture is
Álvarez-Cordero 1996, Sanaiotti et al. 96-100%; annual rainfall fluctuates
2001, Touchton et al. 2002, Ferrari and between 2,000 to 4,000 mm (Cañadas
Port-Carvalho 2003, Muñiz-López 1983). There is a distinct dry season
2007, Piana 2007, Rotenberg et al. 2012 from December to March, a rainy
and Aguiar-Silva et al. 2014). season from April to July and an
Here I present the first study of the intermediate season from August to
Harpy eagle trophic ecology of November (Cañadas 1983) but the dry
Ecuador. season does not occur in some years
(Geenen et al. 2000). Biodiversity level
Study area is high in the Reserve, including around
490 species of birds, 165 of mammals
This study was conducted in two (10 of them are primates), and around
different regions, one at the east and one 470 plant species per hectare (Geenen et
at the west of Ecuador. The Cuyabeno al. 2000).
Faunistic Reserve is located in the At the western region of the country
eastern region of the country and it the study was concentrated in the
belongs to Sucumbíos and Orellana Esmeraldas Province. This province lies
provinces, close to Colombia and Peru in the Chocó biogeographic region
borders. This reserve has which includes western Ecuador,
590,112 hectares (Ministerio del southeast of Panama, western Colombia
Ambiente 2012) that means about 1% of and Ecuador down to the northwest of
the country’s land area. Five indigenous Peru (Critical Ecosystem Partership
groups live and use natural resources for Fund 2001). Relative humidity is more
hunting and cultivating familiar than 90% on average, mean temperature
properties, but overall the human ranges 23 to 25°C and mean annual
density is low (0.002 inhabitans per rainfall varies between 3,500 to
hectare; ICCA Consortium 2010). The 4,000 mm/year (Jahn 2011). Annual
area is dominated by primary lowland rainfall is unimodal, with a dry season
tropical moist forest between 250 and from May to November and a rainy
326 m a. s. l. (Cañadas 1983), and season from December to April (Neil
flooded tropical evergreen forests (Stotz and Möller-Jorgensen 1999). The study
et al. 1996, Cerón et al. 1999, Palacios area belongs to the "Cayapas-Santiago-
82
Wimbi" Important Bird Area that The number and identification of

encompasses 60,000 hectares of humid items in nests were known using both
and very humid tropical forest (BirdLife direct observation of prey delivery by
International 2015). It is calculated that adults at the nest to feed the young in
this area contains about 6,000 species of combination with analyses of prey
vascular plants, 830 of birds and remnants and regurgitated pellets
142 mammal species (Critical dropped under the nest tree to confirm
Ecosystem Partnership Fund 2001). It or complete the observation data. In one
also includes afro-ecuadorian case, the species were identified when
communities that in our study area have uneaten parts of the prey were collected
0.03 inhabitants per hectare (Minda from the nest platform in nest 1.
Batallas 2002). Climbing equipment was used to reach
the nest tree.
The monitoring period for each nest
Methodology: was the following: For nest 1: 52 days,
Nest 2: 156 days; Nest 3: 87 days, Nest
Prey delivered to seven Harpy eagle 4: 84 days, Nest 5: 77 days; Nest 6: 15
nests from 2002 to 2010 were analyzed. days; Nest 7: 77 days; in total 548
One nest was im the western region of observation days. Data from nest
the country (nest 3) and the rest were in number 6 were only used to complete
the eastern region. They were occupied the spectrum of prey species since it
with one juvenile each that were provided very little information as it
different ages. The age of the juveniles was monitored during a short period of
occupying each nest was based in the time.
first visit according to their plumage Direct observations were conducted
pattern (Fowler and Cope 1964, using 10x40 binoculars from 07:00 to
Ferguson-Lees et al. 2001) and behavior 16:30 H from the ground or from a
(Rettig 1978). To facilitate the analysis camouflaged 28 m observation tower,
of data we divided the age of the 34-50 m from the nest tree.
juveniles in two classes of ages Days between consecutive prey
depending on their development status: deliveries were calculated only when
one clase from 0 to ≤3 months old observations happened during
(nestling) and the other >3 months old continuous periods of ≥ 10 d. Prey
(fledgling). species and hour of prey delivery
83
attempt were recorded. Prey were and equally distributed community and
individually identified to genus or lower values represent less diverse
species using a field guide (Emmons community. A value of 0 would
and Feer 1990) when the adult left them represent a community with just one
on the nest platform or when the species. It is explained by the formula:
juvenile was feeding on it. H Shannon/corr = ∑i (pi lnpi)ln N
Prey bones and pellets dropped on where pi is the frequency of the ith
the ground were collected during each element and N is the size of the sample.
visit to the nest area and saved in zipped Statistical differences in the values
plastic bags. The date, number of bones of diversity between pairs of nests were
and nest location was noted on each evaluated by the procedure proposed by
plastic bag. After that they were Solow (1993) which consists of the
identified to genus or species level by randomization of combined data of
comparison to the remains of animals pairs, followed by the calculation of the
hunted by people of the community near difference in the value of diversity
the nest and comparing bones to a between the pairs of samples and the
reference animal collection at the repetition of the procedure 10000 times
Ecuadorian Museum of Natural from which it is estimated the
Sciences in Quito, Ecuador. significance (p).
Mann-Whitney U Test was used to Niche breath was calculated by
determine the existence of any bias in standardized Levins index (Colwell and
regard to the probability of finding more Futuyma 1971):
remnants of some species than others Bsta= (B-1)/(n-1)
under the nest trees. where B is the Levins index
Prey diversity was estimated using (B= 1/∑ p2j), pj is the proportion of
the Shannon-Weaver function occurrence of each prey species and n
(Shannon-Weaver 1949) and Sheldon the number of prey species.
correction (Sheldon 1969) for variable Standardized Levins index values range
sample sizes. This index indicates the between 0 (minimum niche breadth and,
average degree of uncertainty in consequently, maximum selectivity) and
predicting species to which an 1 (maximum niche breadth, minimum
individual belong if it is chosen at selectivity; Krebs 1999).
random from a sample. A high value of Prey weight were estimated based
H would be a representative of a diverse on values published in previous
84
literature (Oliveira 1998, Taube et al. bibliographic references (Pinto et al.

1999, Kays 2000, Poloskey 2000, 2015, BirdLife International 2013)
Robinson and Bennet 2000, Trovati et The conservation and threat status
al. 2010, Catania 2011, Tirira 2011, of the prey was taken into account. Prey
BirdLife International 2013, Pinto and were classified by degree of
Nicolalde 2015). Only adult weights conservation threat in national and
were available for all the species, international categories and were used
consequently only adult prey types were to consider the possible linked pressure
considered for this purpose. on their predator.
To facilitate comparisons of prey
contribution, species were classified
into six different categories: sloths, Results
large monkeys, medium and small
monkeys, other mammals, birds and Prey of seven nests were analyzed.
reptiles. Different development stages of the
Intervals of prey delivery hour were juvenile were found for each nest:
distributed in three categories: 07:00 to juvenile of nest 1 was monitored from 9
10:00 H; >10:00 to 13:00 H and > 13:00 to 12 months old juvenile, juvenile of
to 16:00 H. nest 2 from 6 to 12 months, in nest 3 it
To allocate the prey to a specific was monitored from 15 to 18 months, in
forest stratum the forest was classified nest 4 from 8 to 11 months, in nest 5
based on height classes or elevation from 2 to 4 months, in nest 6 from 18 to
above the ground to select three 19 months and in nest 7 from 1 to 4
different layers: months.
Layer /stratum 1 (low): height from For this study 158 prey records
0 to ≤ 12 m; were analyzed. Diet included 18
Layer/stratum 2 (medium): height identified species (Table 1).
from > 12 to ≤ 24 m; Additionally, six individuals were
Layer/stratum 3 (high): height from recognized within the group of birds,
> 24 m. and fourteen more items could not be
The tree canopy in our study area identified.
reaches 30 to 40 m in height Nests 1, 2 and 5 were those that
(Sierra et al. 1999). The species were contain the highest diversity index
catalogued in different levels in order to (Table 2), and no significant differences
85
were found between them (H´1<H´2; seniculus, 8.9%)) were the most
P=0.85; H´1>H´5; P= 0.93 y H´2>H´5; common species registered. However
P=0.81). the highest contribution of prey in the
Niche breath was Bsta= 0.28 that is a nest 1 was the Order Primate (Kruskall-
low value which shows an Wallis HNest1=5.48; df=1, P= 0.019) and
especialization of the diet of the harpy for nests 2, 3 and 7 it was the Order
eagle. Pilosa (HNest2= 5.18, df=1, P= 0.023;
The sample was clearly dominated HNest3= 8.16, df= 1, P= 0.04; HNest7=
by mammals which comprised 85.2% of 6.1, df= 1, P= 0.014).
prey numbers. There were no significant
Related with prey contribution it differences between species group of
was found that in nests 1 and 2 there prey delivered by adults and
were a bias toward the probability of development stage of the juvenile in the
finding more remnants of sloths than nest (Pearson Chi-square= 9.82; df= 5;
other prey species under the nest trees, P=0.081) and sloths and primates were
so we decided to take into account only the principal type of prey for both
the observation records of prey delivery nestlings (87.8 %, n= 41) and fledglings
to the nest (See table 4). Table 3 shows (90.4 %, n= 94).
filtered prey records. Data on prey left on the nest
In general and taking into account platform were collected only once, in
the taxonomic level (Order and September 2002 at nest 1. The platform
Species), the Order Pilosa and the Two- was composed of a mixture of
toed sloth species (Choloepus unidentified old bones and sticks on a
didactylus at the east of the country and Kapok tree (Ceiba pentandra). A young
C. hoffmanii at the west side) were who half-bodied Three–toed sloth that was
contributed in a highest percentage to delivered one day before weighed at the
the diet of the Harpy eagle (Order time 300 g and its length was 19 cm
Pilosa: 49,6%; from which Choloepus from the head to the thorax. The lower
spp. contributed with 43.1% and Three- extremities had already been consumed.
toed sloth (Bradypus variegatus) The other prey on the platform was
contributed with 6.5%), followed by a Blue and Yellow Macaw (Ara
Order Primate (39.8%) where Squirrel ararauna), that was delivered the same
monkey (Saimiri macrodon, 14.6%) and day of the collection. The juvenile did
Red howler monkey (Alouatta not eat it until measurements were done.
86
It weighed 1,400 g and measured 45 cm 1.7; n= 63). Related with juvenile

from its cervical vertebras to the end of development, adults delivered a prey
the synsacrum or the beginning of the item every 3.3 days (± 1.4; n = 15)
caudal vertebras. It was decapitated by when the juvenile was ≤ 3 months old,
the adult. and every 3.9 days (± 1.8; n= 53) when
Mean weight of delivered prey it was > 3 months. There were no
species was 4.01 kg (range 0.5 - 9.8 kg, significant differences in delivery rates
S.D.= 2.8, n= 44; Graph 1) and there comparing juveniles with different
were no significant differences between development degree (nestling or
the averaged weights of species fledgling) (U=318, P = 0.2; n = 68).
delivered in different nests (Chi-Square Adults delivered prey preferably
=3.97, df= 10; P= 0.9). In addition, the during the morning from 07:00 to 10:00
weight of prey species provided by H (Kruskal-Wallis H= 12.6, df=2; P=
adults to the nest is similar regardless to 0.002; n= 103), and the main prey
the juvenile age (nestling or fledgling) (sloths) were delivered preferably
(Mann-Whitney U= 1668.5, P = 0.15) during the morning (U = 437; P =
when they are grouped in light-medium 0.001). Males were who contributed
weights (≤ 4 kg) and heavy prey (> 4 more to the prey delivery (74.8% n=
kg). 103; Wilcoson Signed Ranks Test = -
2.201; P = 0.03). Contribution of males
and females was maintained for each
juvenile age group (Chi-Square= 3.47;
df=1; P = 0.06).
Sloth prey deliveries were negatively
correlated to primate deliveries (Pearson
Correlation Coefficient= -0.86; P=
0.05). A lineal regression was
conducted showing that sloths prey
Graph 1: Distribution and dispersal of the delivery influences in a 66.1% primates
Harpy Eagle species prey weights in each of
deliveries. Increasing sloths prey
the nests.
deliveries lead to lower primate
comsumption and viceverse (Adjusted
On average, adults delivered one
R2= 0.66; p= 0.05).
prey every 3.8 days (range 1- 7; S.D.=
87
The model will fit as: depending on prey community

Y= 68.9 +(-0.72)x estructure, abundance and availability
Where Y = sloths prey deliveries and x= of those. Diet would be supplemented
primate prey deliveries. with prey that would be hunted
The forest stratum that was more opportunistically.
commonly used to hunt by the Harpy Niche breath was higher for our
eagle was the highest level study than for central Amazonian in
(Level/stratum 3) (Kruskall Wallis Test; Brazil (Bsta= 0.17) (Aguiar-Silva 2014)
H= 17.3, P = 0.004, n = 139) although specialization is evident for
Combining the prey data from all both. In our study area all nests were
the nests, all the species delivered located in the influence area for
except two were found under some community hunting activities. We might
degree of threat or conservation expect that opportunistic behaviour for
pressure at a national level and all but hunting is greater in areas where there is
one at an international level (See Table greater pressure on the preferred Harpy
1). eagle prey as monkeys are.
Species contribution:
Discussion
As in the majority of previous
Niche breath: provisioning behaviour reports, we
found that Two-toed Sloth was always
Existing Harpy eagle reports part of Harpy eagle diet and the most
indicate that it preys on a considerable common prey (Rettig 1978 and 1995,
variety of species, most of which are Álvarez-Cordero 1996, Galetti and
arboreal mammals (Rettig 1978, Carvalho 2000. Sanaiotti et al. 2001,
Álvarez-Cordero 1996, Aparicio 2003, Touchton et al. 2002, Piana 2007,
Piana 2007, Muñiz-López 2008, Aguiar-Silva 2014).
Aguiar-Silva 2014). Although in our Something different happened in
study we found a diversity of prey for Belize where the most frequent food
all nests, values of diversity index were items were Common opposum
relatively low. These data make us (Didelphis marsupialis) and White-
suspect the existence of preferred prey nosed coatimundi (Nasua narica), and
and the use of replacement prey no sloths were found. In this study
88
authors argued that the reason of this could lead to an eagle underexploitation
difference with other Harpy eagle prey of monkeys in territories where humans
reports could be that they were breeding and eagles compete for same prey. Only
at the fringe of sloths natural range for the area around nest 1 the
where prey choice and availability may indigenous prohibit to hunt “animals
be limited. In addition, Belize only with hands”, which includes monkeys.
supports two primate species This situation is reflected in the
(Rotenberg et al. 2012). These data proportion of big monkeys captured by
suggest that in case there is sufficient the eagles that is higher for nest 1 (see
density of sloths these would be Table 3).
selected by the Harpy eagle as the main Related with this we could
part of their diet and it would be able to hypothesize that if sloths are the
respond to changes in the availability preferred prey delivered on Harpy eagle
and abundance of prey across variations nests they should be the most frequent
in their diets. When abundant, the prey provisioned even if monkeys are
Harpy eagle should eat only the most more abundant in the foraging area.
valuable prey type that seems to be However we have found that in the area
sloths and monkeys. Inclusion of other where humans do not hunt monkeys
prey types in the diet should depend on Harpy eagles delivered more primates
the scarcity of preferred prey and than sloths to the nest. That could
abundance of more profitable prey. As suggest that our results fit with the
preferred prey abundance declines, diet predictions made for Schluter (1981)
diversity should increase as happened in about the original optimal diet model:
nest 1, 2 and 5. These nests were Frequencies of alternate prey in the diet
located where indigenous communities may be related not to their own
have their regular hunting routes so densities but inversely with the densities
humans would be competing with the of the preferred prey; when prey are
Harpy eagle for food as indigenous abundant, predators should eat only the
people hunt preferably large size most valuable prey type and as prey
primates (de la Montaña 2013). The abundance declines, diet diversity
persistence of hunting may come along should increase.
with a decrease in the relative Models based on the Optimal
importance of large primates (Zapata Foraging Theory (OFT) (Svanbäck and
2001, de la Montaña 2013) and this Bolnick 2005) established the
89
connection between intrapopulation diet continent, occurring in a diversity of

variation and resource availability. Such environments that includes rainforests
models are useful for our study because (Emmons and Feer 1997) where Harpy
they represent a few examples of the eagle is allocated. Nails and hair were
possible rules that describe how found in a pellet under the tree where
individual niches could vary with nest 7 was located. The few studies on
resource availability. In the competitive the Margay suggest that its diet is
refuge model (CRm), for instance, the mainly composed of arboreal mammals
preferred prey is also the same. For low (de Oliveira et al. 2009) including
and high resource availability this species as Capuchin monkeys (Cebus
model predicts that when resource spp.) (Beebe 1925, Mondolfi 1986) so
availability is high, all the individuals Harpy eagles and Margays could
feed on the preferred prey, but as occasionally overlap their diet causing
resource availability decreases, encounters that could origin an
individuals begin to include alternative opportunistic capture of the eagle on the
prey in their diet in a similar order, as feline.
could happen with large monkeys as
preferred prey and sloths as alternative. Prey weight and provisioning rate:
If resource availability decreases
further, all the prey become scarce, and The Harpy Eagle is known to take
individuals expand their diets and tend a variety of larger-bodied species
to consume all the prey. (>= 2.5 kg) (Barnett et al. 2011) and
Future studies need to evaluate thanks to its massive body size which is
intrapopulation variation in different for males 4.0 - 4.8 kg and for females
ecological scenarios and the relative 7.6 - 9.0 kg (Fowler and Cope 1964,
importance of human hunting on prey Bierregaard 1994) it can prey on species
diversity of Harpy eagles to reveal how that weight more than its own body
individuals adapt to variations in mass, as Wooden monkeys. The peak in
different environmental conditions. food demand is likely to occur in the
Remains of Margay (Leopardus middle stages of brood development
wiedii) were found as a new record for when growth rates are highest and it
Harpy eagle prey species at genus level decreases slightly when broods are
although it has a widespread close to fledgling (Newton 1979). In
distribution throughout the American some raptor studies, greater nestling
90
needs lead to increased provisioning all nest were in the hunting influence
rates (Green and Ydenberg 1994, ratio of an indigenous /afroecuadorian
Jenkins 2000, Masman et al. 1989, community. Rettig (1978) and
Olsen et al. 1998, Tolonen and Seymour et al. (2010) reported fewer
Korpimaki 1994); in other studies, days per delivery as the nestling grew.
parents respond by increasing prey size Rotenberg et al. (2012) observed the
(Newton 1986, Palmer et al. 2004). Our opposite trend in Belize. In our study
results about Harpy eagle provisioning we did not detect any difference in the
preferences may suggest that in our provisioning rates throughout the
study area they could obtain prey that development of the juvenile. However,
exceed the needs of the juvenile sample sizes may have been too low to
throughouth the breeding period as the detect statistical relationships between
mean species prey weight is prey delivery rates and nestling
considerably high during all the months development at an alpha level of 0.05.
covering our study. Eagles probably hunt during the
The frequency with which prey morning or late in the evening and save
items were delivered to the nest (prey prey to leave it on the nest the next
delivery rate) in our observations is morning. In our study prey delivery rate
slightly longer than data collected in is higher for males as happened in nests
other studies. Harpy eagles showed the studied in British Guyana (Rettig 1978)
lowest feeding rates yet found among and Venezuela (Seymour et al. 2010)
raptors (Rettig 1978). On average, our but this is an opposite trend to reports
rate in Ecuador is longer than for British from Belize (Rotenberg et al. 2012) and
Guyana where observed rate is one prey Serra da Bodoquena National Park in
every 2.5–3.5 days (Rettig (1978) and Brazil (Martins Pereira and Salzo 2006)
for Venezuela it is every 2.1–2.4 days where females had a greater role. Both
(Seymour et al. 2010). Those parental care and time-activity budgets
differences could be due to different of male and female raptors may vary as
provisioning tactics as result of result of several biotic (e.g., brood size,
variation in prey availability as a age of adults, food availability,
consequence of influences of their competition with other predators,
seasonal distribution or human hunting hunting skills) and abiotic (e.g., day
pressures that keeps some of the prey length, topography, weather) factors
away from important foraging sites as (Boulet et al. 2001, Palmer et al. 2001).
91
Females are much larger than males, its own breeding characteristics (one
and only periodically join in the juvenile every 3 years), habitat loss and
hunting, mainly during the late nestling hunting pressure (Álvarez-Cordero
and postfledgling period (Newton 1973, 1996), Harpy eagle population stability
Eldegard et al. 2003). We will need is also endangered because the
more data to determine sex contribution resources which they are depending on
to assess a trend in prey delivery rates are threatened as well.
through juvenile development.
Acknowledgments:
Hunting preferences:
I thank SIMBIOE, The National of Bird
Ground and medium-level Trust, EcoFondo and Environmental
vegetation may affect the ability to Ministry of Ecuador for their support.
detect prey and hence may influence the Indigenous communities of Aí / Cofan,
success of particular foraging Siecopai, Kichwa and Shuar
behaviours (Bechard 1982, Janes 1985). nationalities from Sucumbios province
With a wingspan of 176 to 224 cm it as well as afroecuadorian people from
should probably be more difficult to Esmeraldas province allowed us to
maintain the required maneuverability work on their lands and participate
for hunting as in lower strata there are collecting information for this study.
more density of obstacles due to the Special thanks to Irina Muñoz Ron,
thick vegetation. Harpy eagle predatory Eddy García Arana, Andrés Gutiérrez
habits occurring more in the upper Bustamante, Carmen Díez, Marcos
canopy may facilitate the accessibility Mallo, Máximo Sánchez, Enrique de la
to the prey, decrease the risk of collision Montaña, Néstor Quimbita, Alexander
and do more successful any attempt of Blanco and volunteers and assistants for
hunting. their hard work in the field participating
in the Harpy Eagle Conservation
Endangered prey: Program of Ecuador. Thanks to Jose
María Gil-Sánchez for improving article
In this study it was shown that all scheme.
the identified Harpy eagle prey were
suffering some kind of threat or
pressure. That means that, additional to
92
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100
Table 1: Relative abundance, estimated body mass, habits and conservation status of prey delivered on seven Harpy eagle nests from 2002 to 2010 in Ecuador.
National International
Estimated body Conservation Conservation
Order Family Species Habits Forest stratum
mass (kg) Status (Red List Status (Red List
Ecuador) IUCN)
Leopardus wiedii 6.0 Crepuscular and All Near Threatened Near Threatened
Felidae
nocturnal
Carnivora Nasua nasua 5.0 Diurnal Low and medium Least Concern Least Concern
Procyonidae Crepuscular and
Potos flavus 3.0 High Least Concern Least Concern
nocturnal
Bradypodidae Bradypus variegatus 3.7 Crepuscular High Least Concern Least Concern
Pilosa Choloepus didactylus 6.6 Crepuscular High Least Concern Least Concern
Megalonychidae
Choloepus hoffmanii 6.3 Crepuscular High Least Concern Data defficient
Alouatta seniculus 8.0 Diurnal Medium and high Least Concern Least Concern
Atelidae
Lagothrix lagotricha 9.8 Diurnal Medium Vulnerable Vulnerable
Callithricidae Saguinus nigricollis 0.5 Diurnal Low and medium Least Concern Least Concern
Primates Cebus yuracus 3.2 Diurnal All Least Concern Vulnerable
Cebidae
Saimiri macrodon 1.1 Diurnal Low and medium Least Concern Least Concern
Callicebus lucifer 1.3 Diurnal Low and medium Least Concern Least Concern
Pitheciidae
Pithecia milleri 2.5 Diurnal Medium and high Least Concern Data Defficient
Scoirodae Sciurus spp. 0.5 Diurnal Medium and high Least Concern Least Concern
Rodentia
Erethizontidae Coendou bicolor 4.5 Crepuscular High Least Concern Least Concern
Psittaciformes Psittacidae Ara ararauna 1.4 Diurnal High Least Concern Not categorized
Galliformes Cracidae Pipile cumanensis 1.4 Diurnal Medium and high Least Concern Not categorized
Squamata Iguanidae Iguana iguana 6.0 Diurnal Medium Not evaluated Least Concern
Table 2: Shannon´s diversity index for species of prey found in each nest. Nests 1, 2 and 5 shows higher
diversity values.
Nest 1 Nest 2 Nest 3 Nest 4 Nest 5 Nest 6 Nest 7

(n=25) (n=37) (n= 21) (n= 7) (n= 23) (n=2) (n= 29)
H´ SD H´ SD H´ SD H´ SD H´ SD H´ SD H´ SD
1.93 0.17 1.99 0.17 1.23 0.22 1.15 0.32 1.91 0.25 0 0 1.14 0.14
Table 3: Relative abundance of Harpy eagle prey species delivered in each of the nests after bias
correction for nests 1 and 2.
Species delivered Nest 1 Nest 2 Nest 3 Nest 4 Nest 5 Nest 6 Nest 7
Leopardus wiedii - - - - - - 1
Nasua nasua - - - - 2 - -
Potos flavus 2 1 - - - - -
Bradypus variegatus 2 2 3 - 1 - -
Choloepus didactylus 1 11 - 4 10 2 12
Choloepus hoffmanii - - 13 - - - -
Alouatta seniculus 5 3 - - 1 - 2
Lagothrix lagotricha 2 1 - - 1 - -
Saguinus nigricollis 1 3 - - - - -
Cebus yuracus 1 3 - 1 2 - -
Cebus spp - - 1 - - - -
Saimiri macrodon - 3 - 1 - - 13
Callicebus lucifer - 1 - 1 1 - -
Pithecia milleri - 1 - - - - -
Sciurus spp. - - - - - - -
Coendou bicolor - - - - 2 - -
Ara ararauna 1 - - - - - -
Pipile cumanensis - - - - - - -
Iguana iguana - - 1 - - - -
Not identified bird - - 2 - 1 - 1
Not identified
- 2 1 1 - 1 1
mammal
Table 4: Mann-Whitney U Test to assess the biass in the probability of presence of each prey class depending on the method of data collection (Pr= Probability collecting rests
under nests; Pob= Probability using direct observation of prey delivery) . The table shows significative P-values (*) in Nest 1 (for sloths and large monkeys) and Nest 2 (for
medium and small monkeys).
Sloths Large monkeys Medium and small Other mammal Birds Reptiles Not identified
Nest
monkeys
Pr Pob
(n) (n) U Pr Pob U Pr Pob U Pr Pob U Pr Pob U Pr Pob U Pr Pob U
55.5 55.5
78 78 81 75
0.6 0.2 p=0. p=0.0 1
1 0.1 0.5 0 0.1 p=0. 0 0.1 p=0. 0.2 0.1 p=0. 0 0 0.2 0 p=0.
(12) (15) 04 3 p=1
19 19 42 1
(*) (*)
90
136 126.5 128.5 124
0.4 0.4 p=0. 1 133
2 p=0. 0.2 0.1 p=0.4 0 0.4 0.1 0 p=0. 0.1 0 p=0. 0 0 0.1 0.1
(9) (31) 038 p=1 0.6
89 9 34 06
(*)
45.5 42 45.5 42 45.5 45.5
0.7 0.7 49
3 p=0. 0 0 0.1 0 p=0. 0 0.1 p=0. 0.1 0.1 p=0. 0 0.1 p=0. 0 0.1 p=0.
(7) (15) p=1
72 15 48 16 48 48
4 - - - - - - - - - - - - - - - - - - - - -
92.5 95.5 92 97.5 90 71.5
0.5 0.4 1
5 p=0. 0.1 0.1 p=0.7 0.2 0.1 p=0. 0.2 0.2 p=0. 0.1 0 p=0. 0 0 0 0.3 p=0.
(11) (18) p=1
73 2 59 92 2 06
0.5 0.5
0.5 1 1 1 1 1
6 1 (1) p=0. 0 0 0 0 0 0 0 0 0 0 0 0.5 p=0.
(2) p=1 p=1 p=1 p=1 p=1
48 48
97.5 105 105 105 105
0.4(1 0.4(1 112.5 1
7 0 0.1 p=0.1 0.5 0.4 p=0. 0.1 0 p=0. 0 0 p=0. 0 0 0.1 0 p=0.
5) 5) p=1 p=1
5 71 32 32 32
2641. 2844. 2682. 2832.
2683 2720 2793
0.51 0.43 5 5 5 5
Total 0.07 0.13 p=0.2 0.18 0.23 p=0. 0.07 0.07 0.09 0.03 0 0.01 0.07 0.1 p=0.
(55) (104) p=0. p=0. p=0. p=0.
4 47 62
36 89 09 46
CAPÍTULO 5
Pollo de águila harpía en su nido sobre un ceibo (Ceiba pentandra) rodeado de un bosque típico de
tierras inundadas de la Amazonía. Foto: P. Oxford /PCAHE-SIMBIOE
Ñanda gi ña’me ñotssia tsampi jin’ttima isu. Tse’ttini tsu osha’cho aña’cho’qque jin, toya’caen
na’en’su’qque.
Seje’pa’qque, ingi semanqque’su’qque. Toya’caen tsu ande’qque sisipa andepa ñotssi tsuipa jacañe’qque,
mingae asapa’chosa’ne. Tsa’cansi gi antte’fayambi cocama tsa andema itsaye’ja.
In’jamba’qque tsendeccu jipa ingi andembe na’suve dasa’ne.
Guillermo Quenamá, indígena Aí / Cofán de Ecuador
105
106
Capítulo 5
HÁBITAT DEL ÁREA DE NIDIFICACIÓN Y ESTIMACIÓN DE LA

DENSIDAD DE NIDOS DEL ÁGUILA HARPÍA EN LA RESERVA
FAUNÍSTICA CUYABENO, ECUADOR.
RUTH MUÑIZ-LÓPEZ y HÉCTOR ALCÁNTARA
Resumen
A través de la información obtenida de 15 nidos de águila harpía (Harpia harpyja)
localizados en la región nororiental de la Amazonía ecuatoriana estimamos la densidad
de lugares de cría utilizando el método de Densidad Máxima de Nidos. La altitud media
de los lugares de nidificación fue de 219 m s. n. m. (± 11.7 m) y todos los nidos se
construyeron sobre árboles emergentes. La distancia mínima entre nidos más cercanos
fue de 4.9 ± 0.7 km y un área de cría de 19.6  5.7 km2 por cada pareja. La densidad de
nidos en nuestra área de estudio se estimó en 5 nidos / 100 km2. La mayoría de las áreas
de cría se localizó en bosques inundables y relativamente lejos de áreas urbanas. El
hábitat potencial del águila incluye bosques al este y oeste de la cordillera andina pero
los bosques de la región occidental poseen un alto grado de alteración por lo que las
condiciones para mantener una población sana de águila harpía en esa zona son más
limitadas. Algunas acciones para conservar el hábitat del águila harpía incluyen
programas de educación y sensibilización, integración de la población local en los
planes de conservación y desarrollo y voluntad política para controlar la explotación de
los recursos naturales.
Palabras clave: Águila harpía, área de cría, densidad, Ecuador, hábitat.
107
HARPY EAGLE BREEDING HABITAT AND NESTING DENSITY

ESTIMATION IN CUYABENO WILDLIFE RESERVE, ECUADOR.
RUTH MUÑIZ-LÓPEZ y HÉCTOR ALCÁNTARA
Capsule: Using the information obtained from 15 Harpy eagle (Harpia harpyja) nests
located in the Northeastern region of Amazonian Ecuador we estimated the density of
breeding areas applying the Maximum Packed Nest Density method. Altitude of the
nesting sites averaged 219 m.a.s.l. (± 11.7 m) and all nests were built on emergent trees.
Nearest-neighbour distances averaged 4.9 ± 0.7 km and an area of 19.6  5.7 km2 per
breeding pair. Estimated nest densities was 5 nests/ 100 km2 in our study area. There is
a regular distribution of breeding territories which have an extension of 19.6 ± 5.7 km2.
Most nesting areas were located in Flooded forests and relatively far from urban areas.
Potencial habitat includes forests at east and west of the Andean mountains but Western
Region forests are highly disturbated and the conditions to sustain a healthy Harpy eagle
population are more restricted than in the Eastern region. Some Harpy eagle habitat
conservation actions include educational and awareness programs, integration of local
communities in conservation and development plans and political will to control natural
resources exploitation.
Key words: Harpy eagle, breeding area, density, Ecuador, habitat.
108
Introduction species extinction (Brashares et al.

2001, Parks and Harcourt 2002,
South America has the highest Woodroffe and Ginsberg 1998).
absolute number of living species As fragmentation rates continue to
among the seven continents, by a rise in conjunction with expanding
substantial margin (Stotz et al. 1996). human populations (Bierregaard et al.
The major waves of extinctions in the 1992, Laurance and Curran 2008), the
Neotropics are occurring among species fate of tropical rainforests is dubious.
that have evolved ecological The most pressing conservation
specializations that limit their ability to goal today is to reduce these extinction
adapt to human modifications of their waves to a minimum. Effective
habitats (Stotz et al. 1996). conservation of species requires
For many tropical raptors, habitat information about the location of
loss and fragmentation have contributed suitable habitats and the environmental
to population declines (Thiollay 1993, factors affecting them (Guisan and
Bildstein et al. 1998). Rainforest Thuiller 2005). Despite of that our level
fragmentation occurs when habitat loss or knowledge about species distribution
results in an area of continuous forest and abundance in Neotropics is still
being dissected into smaller, isolated rudimentary for all but a few groups of
areas, or ‘fragments’; simultaneously animals and plants.
increasing the amount of forest edge Birds of prey are notoriously
whilst reducing forest area (Broadbent difficult to survey in tropical forests,
et al. 2008, Ewers and Didham 2005, especially in tall, dense, large unbroken
Ranta et al. 1998). Edge effects can tracts of humid lowland forests. In
strongly influence the structure, addition, ecological theory predicts that
ecology, species composition and tropical top predators as raptors are
biodiversity of a fragment though a occur at low densities (Forsman and
series of effects (Broadbent et al. 2008, Solonen 1984, Thiollay 1989b) and
Laurance et al. 1998, Murcia 1995), study areas of 500 ha or more may
placing many specialized species at contain less than one full territory for
great risk of extinction (Cordeiro and most species (Robinson and Wilcove
Howe 2003). Human population size, 1989). However, many rain forest
areal extent of reserves, and isolation of raptors are now threatened by habitat
ecosystems are strong predictors of destruction, disturbance or
fragmentation (Thiollay 1985). There is contribute to the development of

still an urgent need of basic data on successful conservation programmes
natural distribution and density of rain (López-López et al. 2006, 2007).
forest raptors because of a concern The Harpy eagle is the largest and
about the suitability of many reserves most powerful raptor found in the
which may well prove to be too small Americas (Collar 1989) and one of the
for long term survival of some raptor biggest eagles in the world (Collar
species supported originally (Thiollay 1989, Sick 1997).
1989b). It inhabits tropical and subtropical
Like many birds, raptors are usually lowland rainforests (Stotz et al. 1996).
highly selective with respect to their These large raptors apparently occur in
habitats, especially regarding the relative low density and are difficult to
availability of suitable nesting areas, detect in the dense forest canopy
although foraging habitats may also (Álvarez-Cordero 1996). Although not
have an important effect at the time of particularly shy, the Harpy eagle is very
choosing a site during the breeding secretive and spends long periods
season (Newton 1998). Breeding habitat perched motionless (Thiollay 1989b).
(which include both nesting and It feeds mainly on arboreal mammals,
foraging habitats) may limit species specially sloths and monkeys (Álvarez-
productivity or distribution (e.g. Benton Cordero 1996, Piana 2007, Muñiz-
et al. 2002, Soh et al. 2006). In these López et al. 2007, Aguiar-Silva et al.
cases, increasing availability or 2014).
suitability of preferred habitats (e.g. The Harpy eagle range extends
restoring nesting habitats or increasing from Southern Mexico to Northeastern
the availability of foraging habitats) Argentina between 0 to 800 m a. s. l.
may potentially lead to increasing (Ferguson-Lees and Christie 2001,
population sizes (Carrete et al. 2002, Vargas et al. 2006) but in Ecuador one
Hiraldo et al. 1996). Understanding the pair was reported at 1,100 (Pilataxi and
strength of the relationships between Salagaje 2014 through Jocotoco
habitat and species distribution or Foundation comm. pers) and one
breeding success may be important to individual at 1,650 m a. s. l. (Navarrete
manage protected areas and to predict 2004).
how changes in habitat may influence Harpy eagles have been extirpated
population dynamics, and thus from several locations, and are currently
110
declining in various countries (Vargas opposite to inter-nest distances used in

et al. 2006, BirdLife 2013). It is other countries.
considered that the main reasons of its There is no published studies about
population decline are human cuantitative descriptions of Harpy eagle
persecution and deterioration of its breeding areas features at a landscape
habitat (Ferguson-Lees and Christie scale. Before this study Álvarez-
2001, Vargas et al. 2006, BirdLife Cordero (1996) described the ecological
2013), although there is little and geographical range of the Harpy
information about how Harpy eagles are eagle in Venezuela and Panama
affected by these (Gorzula and Medina showing that it is adapted to a wider
1986, Álvarez-Cordero 1996, Guerrero array of forest environments from
1997, Trapé-Trinca et al. 2007). Lowland Rainforest to Tropical Dry
At a global level it is catalogued as Forests; On the other hand, Giudice
Near Threatened (BirdLife 2013) but as (2005) described forest structure and
Vulnerable for Ecuador (Granizo et al. nest tree architecture for nine Harpy
1997). eagle pairs in Madre de Dios (Peruvian
There are few studies about Harpy Amazon). He concluded that in the
eagle nesting density. Vargas and Terra Firme forests of the lower basin
Vargas (2011) estimated 16-24 km2 per of Tambotata river Harpy eagle nests
pair depending on the method used in were determined by a lower basal
one area of Darien region in Panama coverage (average amount of an area
and extrapolated results from 18 occupied by the cross-section
breeding selected pairs to assess the of tree trunks and tree stems) than in
population size for all the country. They randomly selected areas. After that, a
also roughly described some lanscape study in a region of Panama correlated
characteristics as most nests (n= 30) vegetation structure with presence of
were located below 310 m in primary Harpy eagle nests finding that number
forest. In Peru, Piana (2007) calculated of tree families and average tree height
2
an area of 43 km per pair (n = 3 nests) were the best predictors of nest site
and Álvarez-Cordero (1996) determined selection (Vargas et al. 2014).
45-79 km2 for Venezuela (n= 9 nests). Here we present preliminary spatial
Thiollay 1989 a,b, considered 100 km2 landscape-scale analyses of Harpy eagle
per pair in French Guiana but this was breeding areas and an estimated nest
based on individuals observations density in a region in the northeastern
111
Amazon Basin in Ecuador. Assuming widespread type of vegetation in the

homogeneity it can be useful for country, which covers more than a third
providing a crude description of the of the continental Ecuador (Neill 1999).
distribution patterns without any The percentage of relative humidity is
underlying assumption. Thus we present about 72% (61.8 - 76.7 %) and average
a map of the potential distribution of canopy height reaches 35  9 m
this species in Ecuador. (Rivadeneira-Roura 2007).
Its seasonal variation consist of a
Study area dry season - defined as the months that
receive less than 250 mm of rain per
The study area comprised month and includes from late December
approximately 590,112 hectares and it through March; a rainy season - defined
was composed of some sectors of the as the months receiving more than
Cuyabeno Wildlife Reserva in the 250 mm of rain per month, that runs
northeast of Ecuador, bordering from April to July- and the time of
Colombia and Peru, and the fluctuation ranges from August until
northwestern Reserve boundary that is mid-December (Rivadeneira-Roura
occupied by Secoya indigenous group 2007).
and colonist (Ministerio del Ambiente Cuyabeno Reserve embodies an
2012). The Cuyabeno Wildlife Reserve extraordinary degree of biological
is one of the most important protected richness (Araya and Peters 2000). This
areas in Ecuador (Balslev et al. 1998) Reserve has registered the greatest
comprising about 12% of national density of tree species in the world,
protected land (World Resource 473 species per hectare (Valencia et al.
Institute 2005). Located in the 1994). Some taxa are extremely
northeastern portion of the Ecuadorian biodiverse as well: 500 species of birds,
Amazon, between 250 y 326 m a. s. l., 165 species of mammals, and
it is categorized as a Humid Tropical 135 species of amphibious and reptiles
Forest bioma with an annual mean (Ministerio del Ambiente 2012).
precipitation that ranges from 2093.8 to Cuyabeno Reserve was designed to
2728.2 mm and annual average protect the Cuyabeno Lakes District, an
temperature from 24.6 to 26.3 °C ecosystem with unique hydrologic
(Cañadas 1983, Ministerio del characteristics comprising
Ambiente 2012). This is the most 14 interconnected lakes and seasonally
112
inundated lands located at the The Reserve is also home to several

confluence of the Ecuadorian Amazon’s indigenous communities (see Table 1):
black and white water river systems Siona, Secoya/Sieco´pai, Cofán/ Aí,
(Coello-Hinojosa 1992). It was also Kichwa, and Shuar that all have
planned to be a tool to increase the different histories of migration to the
income of local people by improving area and different degrees of
wildlife management and tourist acculturation and market integration
facilities. (Holt et al. 2004).
Cuyabeno river, with its headwaters
in the Andes, is the main river that Table 1: Indigenous communities population
in the study area.
flows through the area and drains into
Community Indigenous Population
Aguarico river, that flows in the
Name Nationality
southeastern region of the Reserve; it
Puerto Siona 111
joins Napo river at the Peruvian
Bolívar
boundary and eventually flows into the
Amazon River. Tarapuy Siona 100
The forest in our study area has two Playas de Kichwa 242
distinctive subtypes according to local Cuyabeno
hydrology (Valencia et al. 1994): Siecoya Secoya/Sieco´Pai 150
- Flooded forest: Irregularly and Remolino
almost permanently inudated forests Charap Shuar 42
flooded by white and black-water Zábalo Cofán/Aí 71
rivers (Asanza 1985, Pires and Prance Zancudo Kichwa 100

1985). Flooded palm swamps occurrs Poocoya* Secoya/Sieco´ 9
along depressions and stream valleys Pai
with a vegetation dominated by Wajosará* Secoya/Sieco´ 7
Morete palms (Mauritia flexuosa). Pai
- Terra Firme forest: This forest Data from UCODEP (ICCA 2010)
* Muñiz-López R. Pers.obs.
occur on low ridge slopes and crests
that are never flooded, known as Terra
Indigenous communities and
Firme. They are well-drained forests
petroleum development existed in the
located on small hills in the upper
area prior to the designation of the
watershed and the areas between the
Reserve boundaries. The indigenous
semi-inundated planes.
113
groups (e.g. Secoya/Sieco´pai) also confused and made mistakes in

established communities within the recognition of the species.
current Reserve boundary. Colonists Bigger efforts in searching nests were
have established themselves in the area made in the lower Aguarico river as the
as well. PCAHE worked more intensely in
Human activities that take place Aí /Cofán territory.
within the Reserve, therefore, range Finding Harpy eagle nests in the
from oil exploration and tourism to short term is a work that can hardly be
agriculture (Erlien et al. 2005). Annual done without the cooperation of the
rates of deforestation within the local people, because they are the ones
-1
Cuyabeno Reserve was 0.08% /year who best know their own territory.
for 1996-2002 loosing 1,300 ha of We estimated the number of
primary forest during 1986-2002 for territorial pairs per unit area. We based
agricultural, livestock and infrastructure these estimates on the spacing amount
construction (Mena et al. 2006). neighboring pairs, indicated by active
nests.
Methodology Since our study nests were not
generally in predetermined study plots
Nest searches and nesting density: that had been exhaustively searched for
them, we used the average distance
Nests were located by the local between neighboring nests to estimate
population in their hunting or fishing densities of territorial pairs. Nests
routes in the framework of the Harpy position were recorded with a hand-held
eagle Conservation Program in Ecuador GPS and plotted on a digital map.
(PCAHE in spanish) between the years Nearest-neighbour distances
2000 and 2011. The PCAHE team (NNDs) between pairs were used as a
confirmed the species that was nesting measure for the Maximum Packed Nest
and organized training activities to help Density (MPND) method to set an
people identify this species of other upper limit on the density that could be
eagles with crest feathers in the study possible for the available habitat under a
area as Ornate hawk eagle (Spizaetus given mean inter-nest distance. To
ornatus) and Crested eagle (Morphnus calculate distances, each of a group of n
guianensis) as they were sometimes nests is joined by a straight line to its
nearest neighbor, yielding n-1 inter-nest
114
distances and a single mean inter-nest where r is half the mean inter-nest
distance (Gower and Ross 1969; Selas distance, adjusted by multiplying by a
1997). NNDs were calculated using constant that includes the portion of the
known occupied nests. non-overlapping area between
To avoid inflating estimates with neighboring territories (Brown 1975,
sites that were obviously not, or might Whitacre 2012). This equation assumes
not be, ‘nearest’ neighbours, only was that spacing is completely regular and
used adjacent, closest or clearly that the maximum possible packing of
neighbouring sites in contiguous or nesting territories is achieved.
nearly contiguous habitat. We excluded Neighbor nest distances offered a
nests that were separated by large way of estimating overall carrying
expanses of open landscape or which, if capacity in areas of known suitable
within the same block of forest, were so habitat. An estimate of pair spacing can
far apart (arbitrarily defined as 2.5 times be used to extrapolate population
the minimum inter-nest distance as used carrying capacity if the area of suitable
for Philippine eagle (Pithecophaga habitat is known. Carriyng capacity may
jefferyii) distribution and nesting study be an important estimate for setting a
in Mindanao Island (Bueser et al. 2003) target population size for endangered
that they might include other, species.
undiscovered, territories.
GMASD statistic of Brown (1975) Habitat characteristics:
was used to test whether nests were
more evenly distributed than if sites A circular plot (radius = 1/2 NND),
were selected by the birds at random. using the mean NND, was drawn
To do that we take the ratio of the around all nests and pair locations,
squares of the geometric and arithmetic including additional isolated or non-
means of nearest-neighbour distances. neighbouring sites. The extent of each
Values of this ratio greater than 0.65 type of vegetation cover (Flooded
indicate regular spacing, while lower forest, Terra Firme forest and
values indicate randomness (Nilsson et Degradated landscape) was determined
al. 1982). within each circular plot using GIS
The area per nesting pair (A) is software (ArcGIS by ESRI).
given by: Degradated landscape included roads,
2
A= r *1.158 oil palm plantations and residential
115
lands. We also took into account when each of the variables is a perfect
physiographic features such as function of the other (Spearman 1904).
elevation, vegetation cover, distribution Kruskal-Wallis one-way analysis of
of water or human development. variance tested whether samples
Altitude was taken in account and it was originate from the same distribution. A
estimated by a hand-held GPS. significant Kruskal-Wallis test will
Habitat-related terms follows Hall indicate that at least one sample
et al. (1997). In brief, habitat is an area stochastically dominates one other
suitable for an organism to use. Habitat sample (Kruskal and Wallis 1952).
composition in the study area was Factor analysis (Thurstone 1931)
determined using the 1: 250,000 digital using Principal Component Analysis
map of Ecuador geographical features (PCA) with variance maximizing
generated by Ministerio del Ambiente (varimax) rotation was used to detect
del Ecuador and EcoCiencia (2005) and structure in the relationships between
the 1: 1,000,000 vegetation estructure forests subtypes and spatial location of
created by Sierra (1999). the nests. The goal in varimax rotation
Nine variables were taken into is to maximize the variance (variability)
account for analysis: altitude (A), of the "new" variable (factor), while
percentage per circular plot of rivers minimizing the variance around the new
(Rv), Terra Firme forest (TF), Flooded variable.
forest (FL) and degradated lanscape The thematic layers of vegetation
(FD), distance to indigenous settlements types that were located below 1200
(Di), distance to forest edge (Dfe), m.a.s.l. were superposed using GIS
distance to urban areas (Dua), and software (ArcGIS) to create a digital
distance to roads (Dr) (see Table 2) . A map of potential areas in Ecuador where
modified Kolmogorof-Smirnov test was Harpy eagles breeding pairs could be
used for testing normality of the data located.
distribution (Massey 1951). Data were
analyzed using descriptive and non
parametric statistics. Spearman´s
correlation coefficient measured
dependence between variables.
Spearman correlation of +1 or -1 occurs
116
Results nesting density of 5 nests/100 km2 in

continuously suitable nesting habitat.
Nest searches and nesting density Nests were found at different
distances from indigenous communities
Locations of all nests are shown in and average separation was 5.9  4.9
Map 1. From 2002 to 2011 we located km (range 1.3-20.1 km).
17 Harpy eagle nests. One of them was
an inactive alternative nest and one Habitat types
different was an old nest that was not
used by eagles in the study period of Forests (both Flooded and Terra
time. All nests were built on emergent Firme) contained more than 90% (90.2
trees: 16 nests were in Kapok trees  7.3 %, N=15) of the vegetation types
(Ceiba pentandra) and the remaining in used by the Harpy eagle in its breeding
a Cedrelinga tree (Cedrelinga area. Those that were closer to human
catenaeformis). Only active nests (n= settlements (nests 1, 7, 9, 11, 12 and 13)
15) were considered for the following were the nests which contained some
analysis. proportion of degradated forest defined
All but four nests were located in as mosaics of forests at various stages
the Cuyabeno Wildlife Reserve. of degradation and human-induced
Altitude of the nest sites ranged modification characterized by oil palm
from 206 to 244 m a. s. l., averaging plantations and roads (5  2 %). Rivers
219 m a. s. l. ( 11.7 m, N= 15). occupied the rest of the area calculated.
Nearest-neighbour distances A Chi-Square test showed
(NNDs) between closest pairs averaged significative differences in forest type
4.9  0.7 km (n= 6; range 4.2- 5.9; N= contribution to the nests (Chi-square =
6). 25.707; d.f.=2; P= 0.00). Flooded forest
The GMASD statistic value was (67  6.1 %) was the most
0.98 that means a high degree of regular representative type of forest in the
distribution of breeding territories. Harpy eagle breeding area studied,
The resulting circular plots followed by Terra Firme forest type (23
averaged 19.6  5.7 km2 per breeding  5.6%) (see Graph 1).
pair that means a maximum theoretical
117
Six nests were located within 18.6 

2.8 km of urban areas (> 1000
inhabitants).
Correlations between degradated
forest proportion in a breeding area and
distance from the nest to an urban zone
resulted in a negative relationship
(Spearman´s rho= -0.758; P=0.001).
No correlation was found for
degradated forest in a breeding area and
Graph 1: The boxplot displays a significative
distance to an indigenous settlement
different contribution of forest types to
(Spearman´s rho = - 0.172; P = 0.541).
Harpy eagle breeding areas. Flooded Forest
is the most representative vegetation type for
Initial Eigenvalues nesting sites. Note right-skewed data and an
% of Cumulative outlier in Flooded Forest type.
Component Total Variance %

This correlation between the
1 4.946 54.960 54.960
variables is confirmed by the Barlett´s
2 2.245 24.946 79.906
Sphericity test since its value is
3 .763 8.476 88.381
statistically significant (Chi-square =
4 .648 7.201 95.582
246.136; df: 36; P = 0.000 ) so that the
5 .329 3.657 99.239
procedure of analysis of factors applies
6 .046 .511 99.749
for this study.
7 .014 .160 99.909
The distribution of the variance
8 .008 .091 100.000
explained by the components of the
9 .000 .000 100.000
model is showed in Table 3. Two
factors explained 79.9% of the original
Table 3: Distribution of the variance by
data variance. Table 4 allows to
component
compare the relative overruns of each
The factor analysis showed that the variable by factor; the first factor is
nest-site variables included in this study formed by the variables: Distance to
are highly related and therefore can be urban areas (Dua), degradated landscape
reduced as evidenced by the index of (Df) and distance to road (Dr). All these
Kaiser Meyer Olkin (K.M.O.= 0.6). variables saturated in a single factor
118
because they constitute a distinct group plots are representing the vegetation
of variables within the matrix of type used by pairs and 1,200 m.a.s.l. is
correlations. This factor seems to reflect the maximum altitude. Counting both
the constraints of existing anthropic areas at the west and east of the Andean
influence. mountains a potential area of
2
The second factor collects the 77,849.86 km could be suitable for
variables designated as Flooded forest nesting Harpy eagles in Ecuador.
(FL) and Terra Firme forest (TF), which
would come to represent the forest type
around nests (See Table 4).
Component
Variable 1 2
Dfe .986 .121
Dua .982 .144
Dr .957 .215
Figure 1: Component plot in rotated space.
Rv .791 -.289
A -.756 -.257
If the entire areas of the circular
FD -.706 -.162
plots are used as representing the
FL .116 .952
spacing needed by territorial birds
TF .055 -.944
between each other, and these are fitted
Di .318 .687
into Cuyabeno Reserve (5,901.12 km2),
Table 4: Rotated component matrix.
the simple estimates for the total
A component plot of the rotated maximum number of breeding pairs
factors (Figure 1) shows how the drop to 295 pairs. These calculation
variables associated with the same assume that the birds are maximally
factor appear next to each other in the packed into the available habitat,
common factor space. ignoring all "edge" and territory shape
Map 2 displays a map of available considerations.
habitat showing an hypothetical
distribution of Harpy eagle breeding
pairs in Ecuador assuming that the
entire vegetation types of the circular
119
Discussion vegetation type (Sierra 1999). A

landscape level transect for the Harpy
Altitudinal pattern eagle distribution in the eastern portion
of the Venezuelan Guayana shows lands
Altitude data for nest in our study were the eagle was present that
are similar to altitude levels reported for surpasses 1,000 m elevation (Álvarez-
Amazon Basin of Peru (below 202 m a. Cordero 1996).
s. l.; Giudice 2005) and Brazil (below A study in Panama came to the
150 m a.s.l.; Aguiar-Silva 2014). In conclusion that tree height was one of
Center America altitude was reported the factors that determined the nesting
below 310 m a. s. l. (Vargas and Vargas place of the Harpy eagle, resulting in an
2011) and in Argentina below 515 m a. average value of 16.2 m high (Vargas et
s. l. (de Lucca 1996). Taking into al. 2014). In our study area the most
account that the maximum altitude of common tree selected for this species to
our study area is 360 m a. s. l. build the nest is the kapok tree (Ceiba
(Ministerio del Ambiente 2012) we pentandra). The Kapok tree is a heavily
would expect this result in altitudinal buttressed, canopy-emergent tree. It can
range for our nest sites. reach 70 m in height and it is limited by
Parker et al.(1996) assembled a elevation (it does not exist above 1,500
database of ecological and distributional m.a.s.l.) (Dick et al. 2007).
information for Neotropical birds. They There is a significant decline in tree
assigned the Harpy eagle to the Tropical height with increasing elevation in
Lowland Evergreen Forest type, broadly Ecuador (Girardin et al. 2014). Most
characterized as <900 m in elevation published studies report structural and
and >2,000 mm rainfall per year (Stotz functional differences in Tropical
et al. 1996). Although our nest Montane Cloud Forests compared with
locations are included in this Lowland Rainforests, showing an
description, it is necessary to sample increase in stem density, basal area and
forests at higher altitudes in order to soil organic matter depth and a decline
elucidate the presence of Harpy eagle in tree height, leaf area index and tree
breeding areas as might foreseeable for species richness with altitude (Tanner
the presence of a couple at 1980, Raich et al. 1997). All of the
1,100 m a. s. l. in Ecuador, included in studies associate these trends with
Tropical Montane Cloud Forests cooler temperatures, fog, reduced light
120
incidence and higher relative humidity. al. 2008, Hernández-Camacho and

At 1,200 m a. s. l. in Ecuador tree Cooper 1976).
height ranges 10-21 m (Girardin et al. According to these data it can
2014) so this altitude would contain theoretically be said that it should not
trees with height enough to be selected be unusual to find Harpy eagle breeding
for the Harpy eagle if tree architecture is areas at least at altitudes of
appropiate. 1,200 m a. s. l. as they have availability
The distribution patterns of most of food resources and nesting trees if
animal species reveal clear altitudinal habitat is undisturbed.
preferences and limits, which can
usually be related to their thermal Nesting density
physiology. Food webs may be altered
by elevational shifts (Beck et al. 2008). Raptors nests that are located in
Sixty-nine species of mammals, birds relatively continuous habitat are often
and reptiles have been described as part separated from one another by roughly
of the Harpy eagle diet from Central to equal distances in a fairly regular
South America (Fowler and Cope 1964, manner (Brown 1970, Newton et al.
Rettig 1978, Izor 1985, Chebez et al. 1977). Such distributions permit
1990, Álvarez- Cordero 1996, Touchton population density and size estimates
et al. 2002, Muñiz-López et al. 2007, via sampling and various spatial
Piana 2007, Rotenberg et al. 2012, statistics such as nearest neighbour
Aguiar-Silva 2014) but sloths were the distances (e.g. Clark and Evans 1954,
most frequent prey in their diet followed reviews in: Ripley 1985, Krebs 1999).
by primates (Aguiar-Silva 2014, Opposite to the results reported by
Miranda 2015) Thiollay in French Guiana (Thiollay
Sloths genus Choloepus and 1989b) we found a regular distribution
Bradypus ranges from sea level up to of breeding territories. Thiollay
2,400 m a. s. l. (Britton 1941, Ureña et concluded that Harpy eagles have
al. 1986). Although primates as extremely low densities and patchy
capuchins (genus Cebus) or red-howler distributions. This difference in
monkeys (genus Allouata) are mostly regularity of nest spacing may be due to
lowland species both have been the very small sample size (three
recorded up to 2000 m a. s. l. (Berton et observed individuals and no nest) in his
study as he worked in an unbroken
121
rainforest where there should not exists Silva et al. 2012). Thiollay (1989b)
constraints that could affect the regular estimated an average of 100 to 300 km2
distribution of the eagles. per pair (3.1 individuals/100 km2) based
The use of methods based on the on three observed records in French
distance to the nearest nest is a very Guiana. We will not consider this two
helpful approach to the determination of last reports for analysis because
the patterns of distribution of species. uncertainty of data.
At the theoretical level, it is predicted Our results do not differ too much
that the territory size is inversely related from those of Panama found for Vargas
with the abundance of food so that and Vargas but distances are smaller
individuals will establish territorial sizes than observed for Amazonian region in
that contain adequate resources to Peru and for Venezuela.
maintain their energy needs (Dunk and These differences may be due to
Cooper 1994). NND offer a direct way different availability of resources
of estimating overall carrying capacity between particular study areas. When
in areas of known suitable habitat. resources are scarce territories became
In previous studies about Harpy larger (Pyke et al. 1977). Assuming that
eagle nest density it was found that raptor breeding territories are
neighbour nests distances averaged determined, in part, by food availability
3.8 km and density 10-20 km2 per pair (e.g. Marquiss and Newton 1981,
(n= 5; Álvarez-Cordero 1996) and Village 1982) habitats with greater
4.1 km and 16-24 km2 per pair (n=18; productivity should be, then, in a higher
Vargas and Vargas 2011) in Panama density of breeding Harpy eagle events
(Darien region), 6.3 km and 45-79 km2 and a reduction in the size of territory
per pair in Venezuela (n = 5, calculated (Yosef and Grubb 1994). On the other
for sites < 8 km apart considered to be hand, success finding the shortest inter-
nearest neighbors; Álvarez- Cordero nest distance and sample size in
1996), 7.4 km and 43 km2 per pair in different studies might be causing these
Amazonian Peru (n= 3; Piana 2007), discrepancies. In addition, differences in
and 5 km with no density estimation estimates of nesting density may appear
(n=2) in a Brazilian Atlantic Forest as result of different methodological
reserve- one found in 1992 and other in approaches. Álvarez-Cordero (1996),
2010- but it was no clear if both nests Piana (2007) and this study applied
were used by the same pair (Aguiar- Maximum Packed Nest Density method
122
while Vargas and Vargas (2011) and long tail that may influence its
compared this procedure with the foraging behaviour and selection of
polygon method resulting in a higher breeding sites. Although individual and
area per pair with the second technique environmental factors may influence the
(16 km2 vs. 24 km2). Thiollay did not use of space patterns shown by Harpy
use inter-nest distances as he did not eagles, these birds are also apparently
found any Harpy eagle nest in his limited by their own morphological and
survey but he concluded density using phylogenetic characteristics.
three observed encounters. Large species have high energetic
Our estimates are similar to requirements and presumably must
reported in Panama with the Maximum occupy large home ranges to obtain
Packed Nest Density method. sufficient food (Zachariah Peery 2000)
Analogous undisturbed Lowland but tropical species have lower
Evergreen Rainforest were reported for territorial requirements than those of
both study areas. A homogeneous temperate regions (Keran 1978). For
distribution of resources both trophic example, Crowned eagle
and suitable places for nesting, lack of (Stephanoaetus coronatus) is the only
competitive interactions between large eagle in sub-Saharan Africa found
sympatric species with similar in primary forest. It has similar skeletal
ecological demands (Begon et al. 1996) morphology to the Harpy eagle (del
and the absence of human habitat Hoyo et al. 1994) and its NND averages
alterations or persecution could explain 1.8  0.43 km (range 1.1 - 2.5 km) with
a regular and similar distribution in both an estimated density of 6.5 km2 per pair
areas. (Shultz 2002). This distance is even
There seems to be a general pattern smaller than closest pairs of neighbors
in breeding-range size in areas that found for Harpy eagles (2.3 km in
offers enough resources and have Panama; Álvarez-Cordero 1996) and
carrying capacity to maintain this density is higher than all Harpy eagle
density of Harpy eagle pairs suggesting reported data. On the contrary and to
that phylogenetic and behavioural illustrate differences with temperate
factors could be influential in spatial regions, Golden eagle (Aquila
territorial use. Harpy eagle has specific chrysaetos) may serve as an example
morphological features such as a big because it is a Holartic distribution
body size, very robust tarsi, wide wings species that is similar to the previous in
123
terms of size and weight but its mean Habitat:

NND is 9.7  3.8 km and density
estimation is 285.7 km2 per pair for a Studies on the habitat of the Harpy
southeastern region in Spain (Carrete et eagle are still scarce. The presence of
al. 2001). nests in our study area was influenced
However the Philippine eagle by two main elements: anthropic
(Pithecophaga jefferyi) that is other big pressure and structure of the forest.
sized raptor species which lives in Results of habitat analysis in our study
tropical geographical range and indicates that Harpy eagle breeding
occupies forest habitat, has nest areas occurs especially in Flooded
neighbour distances much higher than Forests which are usually originated
Harpy eagles (mean inter-nest distance along river sides. However it has to be
was calculated at 12.6-12.7  0.9 km taken into consideration that this result
with the minimum at 8.3 km and could be biased as most nests were
130 km2 per pair; Bueser et al. 2003). found near the edges of rivers or
This difference may be due to streams, probably due to the ease of
anthropogenic habitat alteration and access they provide to the nesting areas.
disturbance in their breeding habitat Flooded Forests comprise the
(Bueser et al. 2003). As suitable habitat second major vegetation type in the
is reduced due to fragmentation, Amazon (Ferreira 1997). The low-lying
resources became more restricted and topography of the basin and seasonality
eagles will need bigger territories of rainfall inundate these floodplains for
(Zachariah Peery 2000). up to six months of the year, and the
Breeding areas are used annual water level fluctuation of the
continuously by Harpy eagles as rivers can reach 14 m in amplitude
juveniles until a new reproductive cycle (Ferreira 1997). Although Terra Firme
starts (Álvarez-Cordero 1996, Muñiz- is found to be consistently more
López et al. 2012). Thus it would be species-rich than Flooded Forests (e.g.
desirable to regulate disturbing Balslev et al. 1987, Peres 1997, Patton
activities (lodging, overhunting, major et al. 2000, Haugaasen and Peres 2005
infrastructures and urban development, a, b), the aggregate primate density in
etc.) all year round. Terra Firme Forests seems to be
considerably lower than that in the
species-poor Flooded (Haugaasen and
124
Peres 2005 a, b). Consequently, the total countries of tropical West Africa. (Dick
biomass estimated will be much higher et al. 2007). Kapok commonly
in Flooded compared to Terra Firme colonizes riverbanks, but is able to grow
Forest. (Haugaasen and Peres 2005b). in both Floodplain soils and Terra Firme
Many arboreal folivores (which account soils that exist above the average
for more than half the biomass of non- inundation level.
volant mammals in Neotropical forests) One nest was built on a cedrelinga tree
show an abundance gradient increasing (Cedrelinga cateniformis) in a Terra
from Terra Firme to Flooded Forests; Firme area. This is a characteristic tree
species exhibiting this pattern include of the upper canopy that is restricted to
sloths (Peres 1997; Eisenberg and Terra Firme land; however, it is
Thorington Jr. 1973) that are the main possible to find it also in flooded
prey for Harpy eagles (Álvarez-Cordero habitats and on hills in the Peruvian,
1996, Piana 2007, Muñiz-López et al. Colombian, Brazilian and Ecuadorian
2007, Aguiar-Silva et al. 2014). In Amazon and in Suriname (Duke 1981).
addition, Howler monkeys (Alouatta Considering this information habitat
spp.) are the second main prey for this features of Flooded Forests may offer
eagle and their population density is good conditions for nesting trees and
positively correlated with forest food availability for Harpy eagles.
structural heterogeneity, soil fertility, The records for Venezuela (i.e.
and degree of seasonality, all of which Caura River and Orinoco Delta;
are higher for Flooded Forests (Peres Álvarez-Cordero 1996), and some areas
1997). in Amazonian Peru (Piana 2007),
Flooded Forests hold canopy- indicate that Harpy eagles routinely
emergent trees. Fourteen Harpy eagle utilize flooded and river-edge forests; In
pairs selected Kapok tree (Ceiba Peru five of nine nests were found in
pentandra) to establish the nesting Terra Firme Forests (Giudice 2005).
platform. This is a heavily buttressed That study suggested that this
tree than can reach 70 m in height and preference may be due to the fact that
one of the most common tree species from that forests eagles would have a
inhabiting the Flooded Forests. It has a best place to locate potential prey and to
worldwide tropical distribution growing get greater protection against predators.
in the Caribbean, northern Mexico to Opposite to that conclusion our study
northern South America and in many adduces that Flooded Forests can be at
125
least as suitable and advantageous as forests keep resources available for

Terra Firme Forests could be. Harpy eagles and if they do not suffer
eagle may be able to nest or to forage pressures or anthropogenic
from any of the emergent trees that are disturbances, then the presence of
found both in Terra Firme and Flooded communities may not be converted into
Forests but as an specialist raptor it a factor that disadvantage the presence
might prefer areas that offer a greater of eagles. On the other hand,
amount of preferred prey, i.e. greater competition of resources used both for
abundance of certain species and not so humans and eagles (e.g. food as
much greater diversity, so that Flooded primates or timber species as cedrelinga
Forests may be the best option. or kapok trees) may decrease the
Nests were located in areas with availability of nearby feeding areas or
low anthropic pressure and relatively far nesting trees and this could foment the
away from urban areas. It was expected displacement or disappearance of Harpy
to find this type of result in our study as eagle breeding pairs that are sharing
we worked at Cuyabeno Reserve and territories with humans.
boundaries and it is a well conserved Shootings, persecution and other direct
area where degradated forest percentage human impairment of Harpy eagles as
is still small (0.1 %) (Rodríguez and de removal of live eagles from their habitat
Vries 1994, Ministerio del Ambiente could happen the more near the villages
2015). are and may impact on their population
Indigenous communities were stability.
relatively close of the Harpy eagle nests In our study area indigenous
found in this study. In other communities have coexisted since their
investigation in Amazonian Brazil there origin with the eagles. These form part
were found 15 nests averaging 1.5 km of their worldview and the Harpy tend
(0.6-2.5 km) away from local to be respected by their condition of
communities (Aguiar-Silva 2014). mythological beings that represent
This proximity could be a reflection of forces of nature. It is not usual to have
favorable or acceptable conditions news of shooting or any other
keeped in their territories for years that disturbance in their territories but
allowed Harpy eagles to share their colonist do not have the same
breeding areas close to this kind of relationship with this species and
human settlements. If this conserved conflicts have been reported.
126
Educational and awareness programs (Kassen 2002, Marvier et al. 2004,

could bring the local people to a better Östergård and Ehrlén 2005).
appreciation and knowledge about this The fact that habitat degradation
species and help to reduce this kind of should negatively affect specialists is
pressures not only on Harpy eagles but predicted by niche evolution theory: the
on wildlife. more specialist a species, the more
A few nests were located outside negative its spatial response to
the Reserve boundaries and all were landscape fragmentation and
successful. In spite of the fact that the disturbance (Holt and Gomulkiewicz
area in which they were found has a 2004).
higher degree of Earth´s surfice The Harpy eagle is a specialist
transformation (deforestation and raptor (Álvarez-Cordero 1996; Fam and
agricultural extensification) compared Nijman 2011) and this could be an
with lands inside Cuyabeno Reserve the argument to expect this species to be
state of the forest is still in a relatively negatively affected by landscape
good situation and well conserved disturbance as natural selection has
probably due to the low density of favored their development in stable
human population and a non massive environments (Kassen 2002). Empirical
exploitation of natural resources in that findings suggest that the decline of
area. Habitat degradation is not yet specialist species observed worldwide is
evident but the advance of African Palm likely to be related to human-induced
plantations, new and more populated landscape degradation (Devictor et al.
human settlements are arriving and the 2007).
construction of infrastructure to develop Álvarez-Cordero (1996) suggested
the oil industry is increasingly growing that the eagles were quite tolerant of
in the province. human landscapes as long as the forest
Specialist species tend to be located matrix remained. In Venezuela large
in less fragmented and less disturbed portions of the habitat of this raptor
landscapes than generalist (Devictor et reverted to production forests managed
al. 2008). They are expected to benefit by logging companies. In his study area
from environments that are relatively he described a breeding area surrounded
homogeneous whereas ecological by an intensive logging activity where
generalists should benefit from one pair successfully completed
environments that are heterogeneous incubation and hatched a nestling that
127
later fledged. He monitored six nests different degree of particular human

where eagles completed 1-2 breeding activities.
cycles while the immediate habitat was Habitat loss and human persecution
undergoing logging. In an study in were identified as the main threats that
Amazonian Peru there were reported six affects Harpy eagle populations at an
Harpy eagle nests and four were located international level (BirdLife
in a land that was heavily used by the International 2015). Ecuador
local settlers in order to subsistence exemplifies the challenges in balancing
hunting and collection of palm leaves conservation and development, with the
and fruits. Additionally two nests were highest deforestation rate in South
found abandoned in areas that were less America over the last 15 years (Pinter et
than 100 m of roads and surrounded by al. 2015) but it has about 20% of its
activities of non-mechanized forest territory (excluding marine reserves)
extraction and palm exploitation (Piana under some degree of protection
2007). Other study in Panama found 30 (Ministerio del Ambiente 2015), and
nests, 25 were located in primary forest, this situation positions it as one of the
and five were in areas of human use Latin American countries with the most
(agriculture or fallow farmland > 2 ha area under some type of protected status
were agriculture had ceased less than 10 (Elbers 2011).
yr ago) (Vargas and Vargas 2011). The conflict between conservation
It is known that habitat loss affects and development is evident in the
the survival not only of the Harpy eagle Ecuadorian Amazon, where
but other species of raptors that depend deforestation has been closely linked to
on forests (Burnham et al. 1988, Stotz the oil industry (Kimerling 1990, 1993;
et al. 1996) and perhaps this examples Uquillas 1984) and where more oil
only indicate a short to mid-term industry growth is expected due to the
resiliency of the Harpy eagles in the large oil reserves in this region (Energy
face of massive intervention (Álvarez- Information Administration 2005).
Cordero 1996). Long-term evaluation of Zones of conflict within or
this land management and related surrounding the protected areas emerge
breeding success should continue to where there are trade-offs and
determine to what extent the inconsistencies between development
adaptability of these eagles can hold and conservation, especially in cases
where colonist and indigenous residents
128
have competing land uses and needs. original western Ecuadorian forests
The main threat that affects the integrity remain (Cerón et al. 1999).
of the Reserve and boundaries and its The western region has only 5.1 %
biodiversity corresponds to the existing of its territory protected by the
pressure in the buffer zone and the Government througt four state reserves
header of the Cuyabeno river by the use that comprise 101,434 ha. Although this
of natural resources due to extractive region has experience extensive
activities such as oil, timber and transformation of the original
monocultives, in addition to the ecosystems by intensive agriculture
expansion of the agricultural frontier (Sáenz and Onofa 2005) small patches
and colonization (Mena et al. 2006). of natural vegetation still remain in this
Large-scale studies provide greater heavily altered zone and they could be a
understanding of species habitat shelter of species as Harpy eagle but
characteristics by revealing the nature high habitat degradation, small
of upper level constraints. The future of extension of protected forests, and few
Harpy eagle population at the east of the protected areas reduces the chances of
country has a good projection if finding areas with high biodiversity
deforestation processes and land cover conservation feasibility.
changes are brought under control. At An active Harpy eagle nest was
the west of the Andean Mountains the found in 2004 in the northwest of
circumstances are different. Ecuador (Muñiz-López 2005) but the
Unfortunately, the West Ecuadorian nest tree was destroyed five years later
region is one of the places in the world because of the incursion in the nesting
where biodiversity is considered most at area of informal mining activity.
risk (e.g. Myers 1988, Parker and Carr Despite visits of ornithologists to this
1992). This region is greatly threatened forests and the sporadic interviews of
by habitat loss and fragmentation this research team to the local people
caused by rural-urban development, we did not have any report of sightings
agricultural activities, uncontrolled from the year 2006, which suggests that
logging, and an inadequate management couples remnants may have been moved
of natural resources (Sierra 1996). This to the most inaccessible places and that
is an area that has been severely this species has a very low population
deforested and less than 4% of the density that makes them difficult to
discover at the west. Although in
129
Ecuador the Harpy eagle is considered of this endangered species is crucial to

as a vulnerable species, in the coastal many aspects of its conservation (Manly
region it may be considered as Critically et al. 1993, Noss et al. 1997) as both
Endangered (Ridgely and Greenfield nesting and foraging habitats may play
2001). an important role in limiting bird
Human persecution may be reduced population numbers or distribution (e.g.
with educational and awareness Newton 1998).
programs. Giving indigenous peoples Tropical rainforest habitat
legal rights to their ancestral lands degradation and fragmentation are one
seems to be a good way to ensure that of the most pervasive threats to the
substantial areas of tropical rain forest conservation of biological diversity.
survive. Efforts to conserve Harpy eagle habitat
Individual species cannot be may include to link biodiversity
protected effectively outside the conservation in protected areas with
biological context afforded by the local, social and economic development
community they occupy but “Flagship” (Brown 2002, Hulme and Murphree
species concept (“popular charismatic 1999) involving Integrated
species that serve as symbols and Conservation and Development Projects
rallying points to stimulate conservation (ICDPs), community-based
awareness and action”; Heywood 1995) conservation, political will and
is an important public relations device. extractive reserves control that may be
It provides a way of communicating to a of great help to safeguard the survival
wide audience the plight of a local not only of the eagles but the vastness
ecosystem. Using this techique could be biodiversity that refuge the forests
useful to acomplish ecosystem where they live.
conservation (Stotz et al. 1996).
Managing populations largely Aknowledgements
depends on managing or maintaining
habitat (Anderson and Gutzwiller We are grateful to indigenous,
1994). Habitat quality is involved in the colonist and afroecuadorian
regulation of the raptor populations and communities for allowing us to work in
determines the species’ settlement their territories and for providing us
pattern (Newton 1998). Further with their knowledge and hospitality.
investigation of the habitat requirements Special thanks to Lina Santacruz and
130
Enrique de la Montaña who worked Anderson S.H. and K.J. Gutzwiller.

hard in the field. We also want to thank 1994. Habitat evaluation methods. Pp.
the volunteers and assistants who 254–271 In T.A. Bookhout (Ed).
helped in the field work and to Research and management techniques
Ministerio del Ambiente de Ecuador, for wildlife and habitats. The Wildlife
SIMBIOE (Sociedad para la Society. Bethesda, MD, U.S.A.
Investigación y Monitoreo de la
Biodiversidad Ecuatoriana) and Araya I. and H. Peters. 2000.
EcoFondo Foundation for funding this Participación de comunidades locales en
study. This article is part of the PhD la gestión de áreas protegidas y sus
thesis of R. Muñiz-López at the zonas de apoyo: Primeros pasos en la
University of Alicante. formación de un comité de manejo
compartido en la Reserva de Producción
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Table 2: Harpy eagle breeding areas variables in a plot of 4.2 km radius from the nest site.
Nest Nest Nest Nest Nest Nest Nest Nest Nest Nest Nest Nest Nest
Variables Description Code Nest 7 Nest 9
1 2 3 4 5 6 8 10 11 12 13 14 15
Topography Altitude A 243 210 213 216 210 206 220 212 224 211 230 218 244 225 212
Terra Firme
TF 35.3 0 64.7 34.9 33.7 0.02 12.6 10 0 11.2 65.7 16.7 29.1 0 28.4
forest (%)
Degradated
DF 3.4 0 0 0.3 0.1 0 18.4 0 20 3.1 18.4 13.8 2.7 0 0.1
forest (%)
Flooded
FL 59.2 97 30.2 58.9 59 93.9 65.4 84 76.5 78.6 12 68.2 63.4 98.9 64.7
forest (%)
Habitat
Distance to
forest-field Dfe 10.9 77 70.5 75 82 101 19.2 68.6 2.1 52.4 6.3 9.4 17.1 12.9 75.5
edge
Distance to
Dr 6.2 81 58.8 63.7 70.1 88 13.4 50.1 3.9 43.3 1.4 4.9 14.1 13.2 53.3
roads
River area Rv 2.1 3 5.1 5.8 7.2 6.1 3.6 6 3.4 7.1 3.9 1.3 4.8 1.1 6.8
Distance to
indigenous
settlements Di 3.8 14 3.1 6.4 3.5 20.1 4.6 3.6 2.8 3.5 1.9 7.5 6.7 5.2 1.3
(< 1000
Settlements
inhabitants)
Distance to
urban areas DUA 16 68.5 67.3 74 81 94.6 20.4 65.5 22.9 56.1 16.4 17 20.9 28.6 68.5
(km)
Min Inter-
nest
Ethological DIN 8.2 5.7 3.6 4.8 8.4 18.2 5.9 4.9 8.9 10.7 9.1 20.1 8.2 8.9 3.6
distance
(km)
Map 1: Harpy Eagle nest location and Cuyabeno Reserve forest types.
Map 2: Potential breeding habitat of Harpy Eagle in Ecuador in dark green colour. Note light green points
showing nest locations.
147
CAPÍTULO 6
Macho de Águila Harpía muerto por disparo cerca de la comunidad de Dureno. Sucumbíos. Ecuador.
Foto: F. Solórzano.
Historia de los Shiwiar de Pastaza:

“El día en el que Ramón Gualinga iba a pasar la prueba para superar la transición de joven a adulto, su
tío tuvo un sueño premonitorio. En él visualizó un águila harpía que se le aparecería en el camino y que
se lanzaría hacia él atacándole. En ese momento, Ramón no se podría amedrentar, por lo que allí mismo
tendría que matar al águila con su cerbatana. Ramón salió al bosque y sucedió todo tal y como le había
dicho su tío. Ocurrió el encuentro con el águila y éste la mató con una flecha. Ésa era la prueba que
tenía que superar. Conservó los huesos en una pequeña bolsa de piel y las guardó para que le
acompañaran el resto de su vida. Éstos le darían fuerza, poder y entereza, cualidades que le siguen hasta
hoy día. Salvo en esa ocasión, nunca había matado un águila harpía, pues para ellos encontrarse con
una de ellas en el bosque les confiere suerte para la cacería. Si las matan, esa misma suerte se les vuelve
en su contra”. (Mayo 2001. Entrevista con Ramón Gualinga-dirigente de ONSHIPAE (Organización de
Nacionalidad Shiwiar de Pastaza-Ecuador)

Muñiz R. 2001. Memoria del estudio Biología y conservación del Águila Harpía en Ecuador. AECI (Agencia
Española de Cooperación Internacional. España.)
149
150
Capítulo 6
MORTALIDAD DEL ÁGUILA HARPÍA EN ECUADOR.
RUTH MUÑIZ LÓPEZ
Resumen
Como resultado del monitoreo continuado de la actividad en los nidos de Águila Harpía
en Ecuador se obtuvo información que documenta los eventos de muerte de individuos
de esta especie y sus causas en el área de estudio. Los registros de mortalidad adulta
fueron menos frecuentes que los de mortalidad juvenil (9.4% mortalidad adulta y 28.6%
mortalidad juvenil) y en todos esos casos las muertes se debieron a persecución humana
(disparos). En los juveniles se encontró que las caídas durante el tiempo de aprendizaje
de vuelo pueden provocar el resultado fatal. Además, se anota un caso de disparo y otro
de muerte tras un comportamiento agresivo de un adulto hacia el juvenil en el momento
en el que éste debía comenzar su dispersión. El que éste debía comenzar su dispersión.
………………………………………………………………………
151
HARPY EAGLE MORTALITY IN ECUADOR.
RUTH MUÑIZ LÓPEZ
Capsule: Information about death incidents of harpy eagle in Ecuador were documented
as a result of prolongued monitoring of their breeding activity. Adult mortality was
recorded less frequently than juvenile mortality (9.4% and 28.6% respectively) and all
adults were killed due to human persecution (shots) for different causes. It was found that
juveniles could die if they fall or lose height from the canopy while learning to fly and
become trapped near the ground. In addition, there was a case of shooting on a juvenile
and another of death after an aggressive behavior of an adult toward the youth at the time
that it should start its dispersión. AbcdefghijklmnñopqrstuvxyzAbc should start its
dispersion.
152
Introduction the harpy eagle (Harpia harpyja) gained

from continuously monitoring their
breeding areas. The harpy eagle is a
Mortality rate is an important
long lived tropical forest raptor that is
parameter for understanding the
considered one of the most powerful
structure of a species’ population and its
birds of prey in the world, as well as
population dynamics. At the individual
one of the largest (Collar 1989; Sick
level, the estimation of this rate is based
1997). It is intermittently found from
mainly on information from ring
southern Mexico to northern Argentina
recoveries and other identification
(Vargas et al. 2006) in tropical and
marking tools such as wing tags, colour
subtropical rainforests (Stotz et al.
rings or satellite and radio tracking
1996). It feeds mainly on arboreal
(Meyburg & Meyburg 2009). In most
mammals, especially sloths and
cases, however these records do not
monkeys (Alvarez Cordero 1996;
serve to detect causes of death.
Galetti & Carvalho Jr. 2000; Piana
In most of the large birds of
2007; Muñiz Lopez et al. 2007; Aguiar
prey, the mortality rate during the first
Silva et al. 2014). Harpy eagles breed
year of life approaches or exceeds 50%,
every 2.5 to 3 yr and the resulting
during the second year it drops to
offspring, usually one per pair, fledges
approximately 38 %, and amounts to
at about five months of age (Rettig
less than 10% after that (Fish &
1978; Alvarez Cordero 1996; Muñiz
Wildlife Service 2004). In general
Lopez 2007) and remains in the natal
mortality is higher for younger birds
area under the parental care for at least
than for adults in both passerine and
two years before dispersing (Rettig
larger and long-lived bird populations
1978, Ruschi 1979, Alvarez Cordero
(Newton 1998).
1996; Muñiz Lopez et al. 2012). The
Generally sources of natural
first active nest of a harpy eagle to be
mortality are rarely reported and are
monitored in Ecuador was found in
poorly known. The information
2002 in the northeast of the country
presented in this work is the first about
(Muñiz Lopez 2005). Its longevity in
the natural and non-natural mortality of
the wild is estimated to be 35 years
153
(Lerner et al. 2009). Ecuador. This zone included 15 of 16

In Ecuador, the species nesting areas that were monitored. This
distribution is restricted to several small reserve comprises 590,112 ha
patches of forest in the northwest of the (Ministerio del Ambiente 2012) of
country and more consistently in the humid tropical forest (Cañadas 1983).
east, where Ecuadorian Amazon Basin Culturally, it is divided into territories
starts (Guerrero 1997, Ridgely and of the indigenous nations of Siona,
Greenfield 2001, Muñiz Lopez 2002). Siecopai, Shuar, Cofan or Ai and
The harpy eagle is currently considered Kichwa, which are located close to the
“Near-Threatened throughout its range main rivers.
(BirdLife International 2013). In The second one belongs to the
Ecuador it is classified as “Vulnerable” coastal region of Esmeraldas province,
(Granizo et al. 1997). in northwest Ecuador, where one
It is thought that the main nesting area was located (number 6; see
reasons for population decline of the Table 1) . This zone lies in the Choco
harpy eagle are human persecution and biogeographic area where the
habitat deterioration (Vargas et al. Afroecuadorian culture is present.
2006; BirdLife International 2013), Our data were collected by
although there is little information about direct observation by members of the
how the species is affected by these Harpy Eagle Conservation Program in
(Gorzula & Medina Cuervo 1986; Ecuador (PCAHE in Spanish) and
Alvarez Cordero 1996; Guerrero 1997; biomonitors which are trained
Ghomeshi et al. 2005; Vargas et al. indigenous people or peasants located
2006; Trape Trinca et al. 2008). near the sixteen nesting areas. All
This paper presents the registration nesting areas were visited at least once a
and causes of death of individual harpy week. Observations were made using
eagles between 2002 and 2011. binoculars from the ground or from
towers 25 to 30 m in height. All dead
Study area, materials and specimens were found during these
methods. monitoring activities in the vicinity of
The study was developed in two their corresponding breeding areas.
diferent areas. The first one was the
Reserve Cuyabeno and its buffer zone
in Sucumbios province in northeast
154
Results youth in the canopy and its fall to a

We monitored 53 individual harpy lower and flooded stratum.
eagles which consisted of 32 adults and In other case, a biomonitor
21 juveniles continuously throughout discovered an eight month old juvenile
the 10 years of study. Nine cases of on the ground that was weak but still
death were detected, which represents alive. A veterinary review detected that
17 % of the total individuals monitored. it was too skinny (pronounced
We found that 9.4 % (n=3) of the breastbone) as to have forces to return
monitored adults died during the study to fly. This individual was recovered
period. As shown in Table 1, all adult and reintegrated to its nest tree
deaths were caused by non natural successfully and we were able to verify
causes (gunshot). On the other hand that the juvenile was alive at least six
28.6% (n= 6) of monitored juveniles months after being placed back in the
died during the study period which is breeding area.
67% of the deceased eagles reported in Starvation and drowning can be
this study (n=9). On five occasions identified as causes of death of juveniles
(records 5, 8, 12, 13 and 16, see Table that fall to the ground and cannot return
1) juveniles fell to the ground and dead. to the canopy.
There is one case (record 12; see Table
1) of a juvenile that was found dead on Discussion
the ground a few meters away from the
nest tree located in a flooded forest with Adult mortality
water level of one meter approximately. In parts of their geographic range,
At the age of its death (30 months) it large predators such as harpy eagles
could fly with skill and was beginning come into conflict with humans. Most
its independence from its parents and its often this consists of harpy eagle
first movements of dispersion (Muñiz predation on livestock (Thirgood et al.
Lopez et al. 2012). Days before, an 2005) and this results in the predator
observer noted one of the adults, being persecuted, trapped, or killed,
presumably a parent, performing with corresponding negative effects on
demonstration flights of attack toward its population (Etheridge et al. 1997).
this juvenile that was perched in a tree Through informal conversations
adjacent to the nest tree. This attack with the local indigenous communities
appears that caused an imbalance of the we collected information about why
155
people wanted to kill the harpy eagles. have attracted attention from the
Their reasons fell into four categories: antiquity to the human being
1- Fear of the species: Many (Reina & Kensinger 1991). In
people in the local communities one case a harpy eagle was shot
believe that harpy eagles will to obtain the its claws as a
attack humans, especially trophy and feathers as a Shaman
children. Because of its large ornament.
size, its attentive gaze and its
reluctance to flee when found by 4- Reprisal for intra-community
humans, it gives the impression conflict: rivalries between
that it is going to pounce. In people in a community can
addition, people compare the sometimes have negative
ease of a harpy eagle catching a consequences for harpy eagles.
large monkeywith that of For example, one of the
capturing a young child. PCAHE biomonitors had an
altercation with another
2- Predation of domestic animals: community member. In
Domestic animals are exposed in retaliation, the community
cleared areas around homes member killed the chick that the
close to forests which makes biomonitor was monitoring.
them easy to prey for harpy
eagles. Because of this, eagles Hunting, or other forms of direct
are shot, which occurred in the human persecution may play an
areas of agricultural frontier in important role in the decline of some
northern Mato Grosso in Brazil raptor populations and it is a significant
(Trape Trinca et al. 2008). In conservation issue worldwide (Raptor
our study we found no livestock Research Foundation 2016). In
remains in prey delivered by Venezuela, Álvarez-Cordero (1996)
adults to nestlings (Muñiz- documented at least 45 harpy eagles
López 2011). shot in a ten year study. Reasons for
shooting varied and included fear,
3- Use of harpy eagle body pleasure hunting, and tribal use as
ornaments: The enormous claws “power medicine” (a case in Panama).
and the long and wide feathers In Brazil, reasons for shooting included
156
fear, lack of knowledge, predation of clarify the degree to which raptor

livestock or fear of livestock predation, predation on livestock is exaggerated in
and revenge against environmentally different communities.
conscious landowner (Trape-Trinca et Large biomass raptors such as the
al. 2007). Thiollay (1984) estimated harpy eagle are particularly sensitive to
that 50 villagers in French Guiana shot hunting pressure (Redford & Robinson
50 raptors per year of multiple species 1987; Silva & Strahl 1991) as it was
(Whitacre 2012). Other tropical eagles, found in this study where all adult
such as the African crowned eagle deaths were directly due to man induced
(Stephanoaetus coronatus) face causes. The importance of human-
persecution over much of their range induced mortality should also be
because of witchcraft, food, ornaments considered in the context of population
and threats to humans (BirdLife 2016, dynamics (Lopez Lopez et al. 2011).
Thomsett 2011). For example, the The main threat to Neotropical bird
African crowned eagle is specifically species is habitat loss (Marzluff &
targeted by hunters in the Ebo forest, Sallabanks 1998), although other
Cameroon (Whytock & Morgan 2010) factors, such as pollution, persecution,
and appears in fetish markets in West and the live-bird trade, have negative
Africa (Nikolaus 2001). Human consequences for conservation for a
conflict due to predation of livestock is number of species (Marzluff &
common and raptors are often shot to Sallabanks 1998). Large biomass
protect farm animals from predation raptors such as the harpy eagle are
(Newton 1979). However, a study on particularly sensitive to hunting
the crowned eagle (Buteogallus pressure (Redford & Robinson 1987,
coronatus) in Argentina found that Silva & Strahl 1991). Governments and
contrary to local beliefs, these eagles NGOs should undertake awareness
rarely preyed on domestic animals campaigns and public education among
(Sarasola et al. 2010). Unfortunately, local populations where this problem is
human persecution is the most especially acute.
important cause of mortality of crowned
eagles in Central Argentina. Although Juvenile mortality:
we also did not find any remains of Although no data are available
livestock in our monitored harpy eagle regarding harpy eagle or other tropical
nests, further research is needed to eagle mortality in the first years of life,
157
our observed 28.6% juvenile mortality is probably a brief period of increase in

is below the 50-83% reported for 13 losses while the young gain experience
European raptor species (Newton 1979) immediately after leaving the nest.
but similar to the 15-29% reported for We were able to verify that on three
Bonelli´s Eagles (Hieraaetus fasciatus) of the days when juveniles fell (records
in Europe (Cadahia et al. 2005). 5, 8 and 13 in Table 1) there were
Except in one case (record 1; see strong gusty winds that are typical of
Table 1) all cases of death of juveniles this season of the year (November-
happened by natural causes. Juveniles December) which could have caused the
who are learning to fly or perfecting instability of these juveniles. If they fell
their flying techniques when they are to the ground or to a lower stratum of
one year old or less perform exploratory the forest, they may not have been able
flights and sometimes land in difficult to propel themselves to a higher perch
positions (see Figure 1) when they try to from which they could achieve enough
grab a new perch (Muñiz Lopez 2007). height to fly again staying trapped under
They must be able to correct these the canopy without the possibility of
positions properly to avoid accidents being fed by their parents as we have
such as falling to the ground. This not detected any adults descending to
behaviour was also reported by feed or take care of the juveniles on the
Álvarez-Cordero (1996) in Venezuela ground. In addition if the nest is located
where a branching harpy eagle nestling in a flooded forest, a juvenile that falls
lost altitude and crashed into the lower to the ground is at risk of drowning or
forest canopy when it was learning to getting wet and with wet feathers they
fly. This juvenile returned to perch on will not be able to fly.
the nest tree within a few hours In other long-lived birds of prey,
however. Álvarez-Cordero described a such as the Iberian imperial eagle
similar event in Panama with a juvenile (Aquila adalberti), 45% of reproductive
continually crashing into the vegetation failures are due tofalls to the ground not
while attempting to fly but finally only of the juveniles, but also of the
returning to its nest. eggs or nest platforms. This happens
Physical capabilities of young birds due to moderately strong winds
increase with a resulting decrease in (Calderon et al. 1987).
mortality rates during the course of Adult raptors such as the Iberian
development (Ricklefs 1969), but there imperial eagle or the tawny eagle
158
(Aquila rapax) perform aggressive studies and to develop conservation

displays to their own offspring, strategies.
(Simmons & Mendelsohn 1993) during In summary, this study revealed that
the period when the parents discontinue the short period during which young
food contributions to the offspring harpy eagles are learning to fly is
which begins the stage of dispersion critical and can be dangerous. All
(Alonso et al. 1987). Although this records for dead adults were by
behaviour is not described for harpy anthropogenic causes, but the majority
eagles belligerent conduct may have of juveniles that were found death in the
caused the death of the juvenile that was vicinity of their nests were due to
being seen attacked by its parents accidents during the post-fledging
(reference 12; Table 1). period. Not taking this into account
The detection of dead harpy eagle could result in inaccuragte estimates of
individuals is extremely difficult, population viability. Detailed studies
especially without identification using more sophisticated methods such
markings or tracking tools so it is easy as telemetry are needed to fill
to underestimate the number of knowledge gaps concerning temporal
mortality events. Comprehensive and spatial patterns of mortality rates. In
monitoring of individuals through direct addition, formal and informal awareness
observation can reveal valuable and education programs are essential to
information including the way deaths mitigate or eliminate the causes of
occur on an individual levelas well as mortality due to hunting and
their causes. misinformation in local communities.
Efforts should be made to monitor
the survival of juveniles in both the
breeding and non-breeding areas, since Aknowledgements
studies focused only on adults may not
detect a decline due to death of I wish to thank to indigenous and
juveniles before they become part of the colonist communities who allowed the
breeding population (Kokko & PCAHE team to work in their
Sutherland 1998; Kenward et al. 2000). territories. I thank Irina Muñoz-Ron,
Understanding the elements that affect Erika Vinueza, Andrea Calispa, Lina
the reproductive success of this species Santacruz, Enrique de la Montaña,
is essential for population dynamics Máximo Sánchez-Cobo and Andrés
159
Ortega for helping with data collection BirdLife International. 2013. Harpia
and assistants and volunteers for field harpyja. The IUCN Red List of
work. I appreciate the improvements in Threatened Species Version 2013;
English usage made by Susan Heath cited 2016 January 2: Available from
through the Association of Field http://dx.doi.org/10.2305/IUCN.UK.201
Ornithologists' program of editorial 3-2.RLTS.T22695998A48128299.
assistance. This study received financial
support from SIMBIOE (Sociedad para BirdLife International. 2016. Species
la Investigación y Monitoreo de la factsheet: Stephanoaetus coronatus.
Biodiversidad Ecuatoriana), Ecuadorian cited 2016 January 2; Available from:
Environmental Ministry and EcoFondo http://www.birdlife.org on 02/01/2016.
Foundation.
Cadahia L., V. Urios and J.J. Negro.
2005. Survival and movements of
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164
Number of
Number
individuals Estimated
Breeding of Year of
Reference to reach Date of age at the Cause of
area name breeding nest Sex
number their death time of death
(Territory) seasons discovery
dispersion death
monitored
period
Shot.
Conflict
December
1 Zábala 2 1 2002 13 months Female within the
2002
human
community.
2 Zancudococha 2 2 2002 - - - -
3 Treiki 1 1 2002 - - - -
Shot in
fear. Eagle
4 Tarapuy 1 0 2003 June 2003 Adult Unknown close to the
vegetable
garden.
November Fall from a
5 Pakuyo 2 1 2003 12 months Male
2010 tree.
6 Junco 1 1 2004 - - - -
Shot
because the
2 de Agosto - eagle
7 1 0 2005 July 2005 Adult Male
Dureno approached
to domestic
animals.
Churuyacu
November Fall from
8 Playas de 2 1 2006 3 months Male
2006 tree
Cuyabeno
9 Tsechueki 2 2 2007 - - - -
10 Cocaya 1 1 2007 - - - -
February Shot for
11 Nueva Vida 1 0 2008 Adult Unknown
2008 trophy
Natural/loss
of balance
October
12 Masakay 1 0 2008 30 months Female due to
2008
parental
display?
November Fall from
13 Charap 1 1 0 2009 13 months Unknown
2009 tree
14 Puñuna 1 1 2010 - - - -
15 Arutam 1 1 2010 - - - -
Enero Fall from
16 Charap 2 1 0 2011 9 months Unknown
2011 tree
Table 1: Harpy eagle breeding territories in chronological order according to their discovery year,
including number of breeding seasons and chicks monitored, and causes of death.
165
Figure 1: A young harpy eagle in a dangerous position practicing its flight habilities. Photo:
PCHAE/SIMBIOE
AGRADECIMIENTOS
A. Tentets, joven de la nacionalidad Achuar del Ecuador y biomonitor del PCAHE

Foto: Enrique de la Montaña / https://vagandomundos.wordpress.com
A las águilas harpías que nos permitieron disfrutar y aprender de ellas, por embaucarnos y llevarnos de las garras
hacia su mundo… por habernos regalado tantas experiencias inolvidables.
Al Atsatábahekukuya por creer en nosotros.
AGRADECIMIENTOS
No puedo empezar unos agradecimientos sin hacer un reconocimiento especial a las

personas que acompañaron mi pasión por seguir adelante con esta experiencia de trabajar con
una especie tan “incómoda” como es el águila harpía. Mis padres y mis tres hermanos
absorbieron cada una de las partes de esta vivencia, no sólo apoyándome y compartiendo
angustias y alegrías, sino también, como mi hermana Teresa (o Kungüimi, como fue apodada en
una comunidad Zápara), acompañándome en el campo. Sé que no es fácil que una hija viva tan
lejos y trabaje en tierras tan recónditas como las de la Cuenca Amazónica pero, a pesar de eso,
siempre alentaron ese fuego interno que veían que me hacía feliz y ese esfuerzo tiene, para mí,
un valor incalculable.
Muchas personas jugaron un papel esencial durante el largo tiempo que pasé en las
comunidades indígenas, afroecuatorianas y de campesinos en Ecuador, pero algunas fueron
fundamentales no sólo ayudándome en el trabajo de campo, sino en cuanto a asegurar mi
bienestar. Llegar y vivir de repente en un medio tan ajeno para una española como la selva
ecuatoriana no es fácil si no se tienen personas que te dan su afecto, te acompañan y te ayudan a
comprender dónde estás, integrándote en todas, toditas las actividades cotidianas…: Mis
compadres Gloria y Pascual, Guadalupe, mi amigo Alberto (“Gengis Kan”), Basilio y Julia
Santi, América, mis otros compadres Beatriz y Aníbal, la familia Mucushigua, Juan Santi…
todos ellos en el territorio Zápara. Ya en tierras Achuar, Domingo Peas, Felipe y Juana, Olimpia,
el fallecido Pincho y Simona, Entsakua y Mupish, mis compadres Tirirut y Soledad, Wananch e
Isolina, Sanchim… Y más al norte, en tierras Aí / Cofán: mis compadres Aníbal y Miriam y
todos sus hijos, Oswaldo, el fallecido Atanasio, Vendi, Elba, Dora, mis otros compadres
Floresto y Juana, mi comadre Macarena, John y Ana, Bolívar, Andrés, Juanito, Pablo, Roberto,
Arturo, Benjamín, Mauricio, Mimi, Carlos… y río arriba, en territorio Shuar, especialmente a
Sandra y Ramiro Jisma, y más aguas arriba aún en el río, ya en territorio Siecopai, a Guillermo y
Marina Payaguaje, Hilario, Sandra, Stalin, Lenin, Verónica y todos los hijos y nietos que forman
la comunidad Wajosará. Fuera de la Amazonía, en la “África” americana, mi querido amigo
Germán alegró con su energía y ocurrencias cada visita que hice a sus tierras y, ya hacia la
Cordillera, bajo las faldas del volcán Cotacachi, Vinicio y su familia me acogieron como una
más. Me siento inmensamente afortunada por haberles tenido a todos a mi lado, ¡y por cada uno
de los nombres con el que me bautizaron mientras compartimos tantos años juntos!
Además de las personas de las comunidades, algunos entrañables compañeros que hoy día
son grandes amigos, estuvieron “guerreando” conmigo en el campo… con todos vosotros tengo
tantas aventuras acumuladas como hojas tiene un ceibo… Irina y Lina que, tras ser voluntarias,
más tarde formaron parte formalmente del PCAHE. Ambas son unas “berracas” en el campo, y
cuentan con la no tan frecuente cualidad de comprender y saber integrarse en el modo de vida de
una comunidad indígena, compartiendo el día a día con alegría y respeto. A este equipo
femenino, hay que añadir al Gorri (Enrique de la Montaña), que llegó a nuestras vidas y a
nuestro proyecto, hoy en día también el suyo, con una energía, ilusión y ganas de hacer bien las
cosas como pocas veces he visto. Además de que las personas de las comunidades le adoran, su
aporte ha sido incuestionable para nosotros y poder contar con su apoyo se ha convertido en un
lujo con el que esperamos seguir contando.
Hay una persona que ocupa un lugar especial en mi gratitud y que seguro ni quiere que le
agradezca nada, ¿verdad Paúl? Paúl Tufiño es una de esas mentes y espíritus inquietos que
apuesta por los “imposibles” y que cree en que las cosas, con ganas y energía, salen adelante. Él
me halló donde nadie me encontraba y con él creció y se fortaleció todo este trabajo con las
harpías. Confió en mí como nadie lo había hecho antes en Ecuador, y me abrió las puertas de
SIMBIOE, de su casa, de su familia y, lo más importante, de su amistad. Con él, Gabi se
involucró con convicción en el periplo aguilero y nos acompañó tanto en el campo como en la
oficina, dejándonos hermosos diseños que daban brillo a cada uno de nuestros esfuerzos por
divulgar lo que creíamos importante en nuestro proyecto. Nuestra amistad y cariño también
creció hasta hoy en día.
Mi admirado Alexander Blanco, hermano venezolano, ¿qué hubiésemos hecho sin ti? Tu
entrega, voluntariedad, tesón, conocimiento, alegría y nobleza nos conmovieron desde el
principio. A pesar de romperte una pierna en uno de nuestros ceibos y de huir “a las bravas” de
la propuesta de casamiento de la hermosa hija de mi compadre Floresto, regresaste una y otra
vez para ser pieza fundamental en las capturas de las águilas. Mucho aprendimos todos de ti, y
mi agradecimiento por tu generosidad siempre estará presente.
En el PCAHE hemos tenido personal y voluntarios que dejaron lo mejor de sí para que este
trabajo saliera adelante. Algunos vivieron experiencias únicas, otros vinieron de lejos y unos
pocos repitieron. Seguro que se me olvida alguno, pero gracias: mi Pitu (Julia) o la “Pila Pila”,
inolvidable en todas las comunidades por las que pasó, David, el Indio (José María), José Luis,
Gusti y las increíbles ilustraciones que crearon nuevos libros, José María Hernández, mis
queridos Néstor, Luis Tonato, Javi Escribano, Andrea Calispa, Erika Vinueza, Adrián Soria,
Edgar Freire que luego trajo el latido del teatro a las comunidades, Gabriela Madrid, Solange
Barahona, mi estimado asistente Eddy García, Mariela Castillo, Jacint, Luis, Maricela, Máximo
Sánchez Cobo, Carmen y Marcos (el primo perdido), Carlos y Lise, Nacho, Piluki, Bego,
Migue, Salva, Santiago Molina, Alex, Sergio, César, Hugh, Itziar, Mª Fernanda, Ana María,
Don Melitón y Zulema, Elenita, Fabián, Víctor Matarranz, Juan Manuel, Tatiana, Salva y su
equipo de Uhkupacha, Fiore y los integrantes de Simurgh…
El Ministerio del Ambiente nos ayudó en muchísimas ocasiones tanto con su personal como
con su equipo logístico. Llegar en canoa a los territorios de las águilas no es tan sencillo y,
cuando lo hemos necesitado, siempre nos han echado una mano. Gracias especialmente a Luis
Borbor, jefe de área de la Reserva Cuyabeno, y todos sus guardaparques y personal: Vilma,
Ramón, Patricio, Florencio, Ángel, Nancy y el fallecido Luis. Gabriela Montoya, desde las
oficinas en Quito y también en el campo, ha sido la responsable de la Estrategia de
Conservación del Águila Harpía en Ecuador y siempre ha respaldado y se ha interesado por
nuestro trabajo, lo que ha sido importantísimo para avanzar en nuestros intereses comunes.
Y a la hora de divulgar y dar a conocer a esta hermosa águila tanto en Ecuador como allende
sus fronteras, mil gracias por mantenerse al pie del cañón en esas jornadas difíciles de campo,
cuando el águila no se asoma, cuando llueve, cuando las cámaras se caen al río, cuando hay una
nueva historia que contar… gracias a la editorial Quercus, a National Geographic, Pete Oxford,
Luis Miguel Domínguez y todo su equipo, Julio García, Manuel Fernández y el grupo de
Sogecable y a los programas “La TV” y “Día a Día” de Ecuador, siempre pendientes de una
nueva noticia “harpiera”.
Quiero agradecer también su apoyo, sobre todo en mis primeros pasos con estas
impresionantes águilas, a Karla Aparicio, quien me aceptó como voluntaria allá en Panamá, a
Eduardo Álvarez por sus consejos y Tania Sanaiotti por compartir.
A Carlos, Ángel y David, por vuestra paciencia y por no ponerme trabas para dar rienda
suelta a mis aventuras.
Víctor Hernández me llevó a Vicente Urios, mi director de tesis, al que agradezco su
dirección, consejos y orientación para llevarla a cabo. Le doy mi gratitud también porque a pesar
del tiempo, de mis ausencias en el campo o mi dispersión en los mil quehaceres y proyectos, de
la dificultad de trabajar en Ecuador y de todas sus tareas, siguió pensando que esta tesis era
posible, y me apoyó no solo moralmente, sino también a conseguir que fuésemos nosotros los
primeros en el mundo que equipamos con un emisor GPS a un águila harpía silvestre, un hito en
la historia de la investigación con esta especie. Gracias Vicente por no abandonar y por darme
esta oportunidad.
Por supuesto, los fondos económicos son esenciales para poder lograr llevar a cabo un
proyecto tan caro – sobre todo a causa de las condiciones logísticas- como el de investigar al
Águila Harpía. Quiero expresar mi gratitud a todos los que decidieron que financiar este
proyecto merecía la pena: SIMBIOE, Fundación Indio-Hilfe con mi estimadísima Mascha al
frente, Fundación Terra natura, Conservation Bird Trust (gracias Jemima!), Optics for the
Tropics, IdeaWild, Neotropical Bird Club, EcoFondo y la Beca Reina Sofía que financió parte
de mis estudios de doctorado en Alicante.
A mis amigos de uno y otro lado del océano, gracias por escucharme y dejar compartir mi
vida y mis historias con vosotros… a mis compañeros de doctorado, Beatriz, Ugo, Jose, Lucía,
Valentina, Raúl, Miguel… fuente de inspiración…
Esta tesis se la dedico a Breogán, quien espero pueda recoger algún día el fruto de nuestro
esfuerzo en tratar de dejar un mundo algo mejor…
Maketei.... Chigatsuafepoenjá
Garras de águila harpía, el ave rapaz más poderosa del planeta. Foto: P. Oxford/PCAHE-SIMBIOE

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BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia harpyja)

Ruth Muñiz López

RUTH MUÑIZ LÓPEZ

BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia

BIOLOGÍA Y CONSERVACIÓN DEL ÁGUILA HARPÍA (Harpia harpyja)

Dª. RUTH MUÑIZ LÓPEZ

Tesis presentada para aspirar al grado de

Programa de Doctorado: “Ciencias Experimentales y Biosanitarias”

DR. VICENTE URIOS MOLINER

En Alicante, junio de 2016.

Ruth Muñiz López Vicente Urios Moliner

Foto de portada: P. Oxford /PCAHE-SIMBIOE

ANEXO FOTOGRÁFICO 173

Extraído del poema “Como puma herido”

1.1.2. Antecedentes en la investigación del águila harpía.

Venezuela y en la provincia de Darién en Panamá (Álvarez-Cordero 1996) no se

La primera vez que se recogieron datos de campo acerca de la distribución del

ejemplares procedían de la localidad de Balzar (provincia de Guayas), de la

(1979) ya hizo referencia a la prácticamente nula existencia de registros de la especie en

Churute, y se desconoce su destino; el segundo individuo formaba parte de la colección

1.1.3. El águila harpía: la especie.

El águila harpía (Harpia harpyja) es el accipítrido de mayor tamaño del continente

Actualmente, se cataloga de forma global como “Casi Amenazada” en todo su

1.1.4. Relación con la cultura y reconocimientos nacionales.

1.2. Justificación de la Tesis Doctoral.

Suramérica contiene aproximadamente 92 especies de aves rapaces diurnas, siendo

inversión de tiempo (Bednarz 2007), lo que ha resultado en un escaso número de

1.3. Objetivos de la Tesis Doctoral.

1.3.1. Caracterizar la dinámica de movimiento de individuos juveniles y establecer

- ¿Cuánto tiempo permanecen los juveniles en el territorio paterno?

- ¿El distanciamiento se produce de forma progresiva o repentina?

1.3.2. Determinar el comportamiento de dispersión juvenil. Contrastar la hipótesis

 ¿A qué edad comienzan los juveniles sus movimientos de dispersión?

1.3.3. Describir la dieta y ecología trófica durante el periodo de cría en distintas

- ¿Cuál es el espectro de especies que caza el águila harpía cuando está

1.3.4. Estimar la densidad de nidos en una región del nororiente de Ecuador

- ¿Cuántas parejas de águila harpía puede contener un área protegida como

1.3.5. Examinar las causas de mortalidad en el norte de la región amazónica de

- ¿Cuál es es la tasa de mortalidad detectada en el área de estudio para

- ¿Qué acciones pueden emprenderse para reducir el impacto y grado de

1.4. Estructura de la Tesis Doctoral:

La presente memoria doctoral se compone de un capítulo introductorio y de

A este capítulo (Capítulo 1), le siguen los Capítulos 2, 3, 4 ,5 y 6 que reproducen

A continuación se describe el contenido de los siguientes capítulos:

En el Capítulo 2 (Muñiz-López, R., R. Limiñana, G.D. Cortés and V. Urios. 2012.

A lo largo del Capítulo 4 (Muñiz-López R. Trophic ecology of the Harpy Eagle

1.5. Área de estudio, resultados generales y discusión general.

Mapa 1: Localización geográfica de Ecuador (área de color marrón) en el continente americano.

La Reserva Cuyabeno, localizada entre las provincias de Sucumbíos y Orellana, es

Detectar individuos muertos de águila harpía es extremadamente complicado,

interés de la población acerca de la importancia de conservar la biodiversidad,

1.6. Conclusiones generales de la Tesis Doctoral.

1. Se describen por primera vez los movimientos de juveniles de águila harpía

2. Se estima en 28 meses la edad en la que un juvenil comienza su dispersión

3. En sus movimientos exploratorios de dispersión, el juvenil recorre una distancia

4. El águila harpía presenta hábitos tróficos especialistas aunque es capaz de predar

6. Existen condiciones adecuadas para que el águila harpía establezca su área de

7. La densidad de áreas de cría de águila harpía en nuestra área de estudio es

9. El éxito de cría de águila harpía puede ocurrir cerca de comunidades humanas