FORKTAIL 29 (2013): 100–106

Bonelli’s Eagle Aquila fasciata renschi in the Lesser
Sundas, Wallacea: distribution, taxonomic status, likely
origins and conservation status
COLIN R. TRAINOR, STEPHEN J. S. DEBUS, JERRY OLSEN, JANETTE A. NORMAN & LES CHRISTIDIS
Records of Bonelli’s Eagle Aquila fasciata renschi on 18 islands in the Lesser Sundas, from Lombok to the Tanimbar islands, in Indonesia andTimor-Leste are reviewed, and its taxonomic status examined. The species is resident on many islands, known to breed on several of thelarger islands and is most abundant on Flores and Timor. It appears to be rare on Lombok and Sumba. There is minimal genetic differentiation
between local subspecies A. f. renschi and nominate A. f. fasciata, suggesting only recent geographic isolation and that it should not be
afforded species status. The species may have been introduced to the Lesser Sundas by traders or colonists in the past. The species’s biology
and ecology are poorly known in Wallacea. It occurs in a wide range of sites from sea level to about 2,000 m, in wooded habitats with apreference for tropical forest. The sparse data on diet show that it feeds on introduced wild junglefowl Gallus sp., but presumably it also
feeds on large-bodied frugivorous pigeons, other birds and small mammals. The two reports of nesting were in May and June–July. The
frequency of records on Flores and Timor suggests that these populations are currently secure, but may be threatened by hunting, captureand deforestation. A predator of village chickens, it is likely to be targeted by local communities. Conservation priorities include distribution
and population density surveys and awareness projects throughout its range.

INTRODUCTION
It has been proposed that the isolated Lesser Sundas population of
Bonelli’s Eagle Aquila fasciata renschi should be regarded as a distinct
species (Thiollay 1994, Ferguson-Lees & Christie 2001), partly
based on biogeography. However, this has not prompted specificfield or taxonomic studies of this taxon. According to recent
evaluations the Wallacean subspecies is found on the Lesser Sunda
islands of Sumbawa, Komodo, Flores, Besar, Timor, Wetar andLuang, and Yamdena in the Tanimbar islands (White & Bruce 1986,
Coates & Bishop 1997, Ferguson-Lees & Christie 2001). The
nominate form fasciata has a wide but fragmented distribution; it isresident, with little or no evidence of migration, in North Africa,
the Iberian peninsula, the Mediterranean, parts of the Arabian
Peninsula, the Middle East, Afghanistan, Pakistan, India anddisjunctly to north Indochina and southern China (Thiollay 1994,
Hernández-Matías et al. 2011). The Lesser Sundas are about 3,000
km from the nearest Asian population in Vietnam (Ferguson-Lees& Christie 2001). In Vietnam, Laos, Thailand and Myanmar there
are very few records and it appears to be sedentary (Duckworth etal. in press).
Although the species has a relatively distinctive appearance
there have been identification problems in the Lesser Sundas. The

type specimen was originally identified by Rensch (1931) as a
Changeable Hawk Eagle Nisaeetus cirrhatus— the Lesser Sunda
population of this species is now known as the Flores Hawk Eagle
N. floris (Gjershaug et al. 2004), and a Wetar island record of‘Changeable Hawk Eagle’ (Hartert 1904) was later identified as
Bonelli’s Eagle (Mees 2006). Bonelli’s Eagle is now in the genus
Aquila rather than Hieraaetus, based on DNA data (Wink & SauerGürth 2004, Helbig et al. 2005, Lerner & Mindell 2005)
demonstrating that Aquila and Hieraaetus as conventionallycircumscribed were paraphyletic.
Recent observations confirming the species’s presence on
Lombok and Sumba, and on several other islands and islets in theLesser Sundas, are reviewed, and substantial populations on at least
Flores and Timor are documented. Records were collected frompublished and unpublished trip reports, bird tour reports and grey
literature, and by canvassing individuals and the Orientalbirding
e-group. The taxonomic status of renschi is assessed using DNA
evidence. Some of the text of this paper was originally submitted
as a book chapter in 2007 (still unpublished), which was split into
two chapters by the editor (Christidis et al. in press, Debus et al. inpress) and is expected to be published in late 2013. This
opportunity is taken to substantially update those manuscripts,
particularly in relation to distribution, ecology and taxonomy.

Figure 1. Map of the Lesser Sundas region, showing the islands mentioned in the text.

Forktail 29 (2013)

Bonelli’s Eagle Aquila fasciata renschi in the Lesser Sundas, Wallacea

Study region

The Lesser Sundas are located in the extreme south-east of Asia,
between continental Java and Bali and the Australo-Papuan region(Figure 1). Lying west to east in a 1,700 km arc, they comprise about
20 large oceanic islands and several hundred small islands and islets.
The inner Banda arc includes young volcanic islands (Monk et al.
1997) from Lombok to Damar in Maluku province and the outer
Banda arc, dominated by sedimentary rocks such as limestone,
extends from Sawu (Sabu), Roti, Timor, Moa and Babar to the
Tanimbar islands. Sumba is considered to be a continental fragment.
Most islands are only a few kilometres apart and would have been
contiguous during the last glacial period 9,000 to 18,000 years ago,thus aiding avian dispersal (Voris 2000), but Sumba, Wetar, theTanimbar islands and Damar are separated from their nearest
neighbours by tens of kilometres. The natural vegetation of theislands is closed-canopy tropical forest (tropical dry to evergreen)and various savannahs, including Eucalyptus, but on many islands,
agriculture has repeatedly changed this to regrowth forest andsavannah woodland (Monk et al. 1997). The main islands have beencleared to varying extents, many now being essentially deforested(Table 1). On large islands, forest fragments are often restricted tosteep mountain slopes and peaks, but on some isolated islands in
the Banda Sea (e.g. Wetar, Romang, Babar and Damar) forest coveris extensive.
Table 1. Area, climate (in relative terms), estimated remaining forest
cover and relative biological survey effort, on Indonesian islands
inhabited by Bonelli’s Eagle (source: CRT unpubl. data, who has visited
all islands mentioned, but has only sailed past Luang Island).
IslandArea (km2)

ClimateLombok4,625

wet (dry on coast) 10

Sumbawa

14,386dry

15

Moderate

Komodo

330

dry

40

Moderate

Flores14,154wet (dry on coast) 15

High

Besar

64

dry

High

Forest cover (%)

75

Survey effort
Moderate

Adonara

509

dry

5

Moderate

Pantar

728

dry

15

Low

Alor

2,864

dry

3

Moderate

Sumba

10,711

dry

12

High

Timor

28,418

dry

10

High

Wetar

2,684

dry

97

Moderate

Atauro

147

dry

10

High

Romang

184

wet

75

Low

Luang

5

wet

25

Moderate

Sermata

103

wet

35

Low

Damar

198

wet

75

Moderate

Yamdena

3,333 (+ satellites c.1,500) wet

75

High

RESULTS
Summary of Bonelli’s Eagle records in the Lesser Sundas

101

Endangered Flores Hawk Eagle (Raptor Conservation Society2011).
Sumbawa: a single historical specimen is the renschi holotype(Stresemann 1932), a male (Mees 2006). Two birds were recorded
by Johnstone et al. (1996), who considered it rare. J. M. Thiollay(in litt. 2007) observed it at four locations (Alas, Reloka, Ropang
valley and Tambora peninsula), where it seemed common in June–
July 2003. One bird was seen at Teluk Saleh in 2009 (V. Dinets inlitt. 2013).
Moyo: a pair recorded in December 1999 (Trainor et al. 2006);
first record for this almost land-locked island.
Sumba: observed at Lake Pambotanjara and Lewa on 14 July 1991
(Dreyer 1993), apparently the first records for the island. One bird
was observed at Lewapaku in October 1998 (Trainor et al. 2006).
It was not observed in four weeks by Olsen & Trost (2007),
suggesting that it is uncommon or rare. None have been recordedby recent bird tour visits to the island. Sumba and Flores are visible
from each other and separated by 45 km of sea, which shouldpresent little barrier to an eagle’s flight. Sumba appears to holdsuitable habitat: a dry limestone island cut by canyons, with
abundant parrot, pigeon and junglefowl prey (JO, pers. obs.).
Komodo: recorded in semi-deciduous forest and savannah(Butchart et al. 1994). At least two observed in June–July 2003 (J.
M. Thiollay in litt. 2007).
Flores: two fairly recent specimens, collected in 1971, identified as
male and female (Mees 2006). Coates & Bishop (1997) considered
it locally moderately common. Fourteen records mostly of pairs,
mainly in hilly and mountainous terrain with cultivation, coconut
plantations, scrub and secondary forest; also on a cultivated andscrubby plain (Verhoeye & King 1990). Recorded in moist and semideciduous forest, thorn scrub and montane forest, from sea level to
2,000 m (Butchart et al. 1994). Two observed in coastal scrub, gallery
forest and grassland (Gibbs 1990). Verhoeye & Holmes (1999)
reviewed about 20 records from cultivated and wooded hills; there
are five additional records for forest and cultivated land (Pilgrim etal. 2000), and eight sightings (including a pair twice) from six
localities in September–November 1998 (R. Drijvers unpubl. data).
Recorded at 12 locations by Trainor & Lesmana (2000). Observed
at eight locations in June–July 2003 and found to be widespread on
the island; one nest found (J. M. Thiollay in litt. 2007). Observed
throughout the island, from sea level to at least 1,600 m in forest,
rice fields and valleys; common in central Flores; a captive juvenile(Plate 1) observed on 14 October 2004 originated from the slopesof Gn Iya, near Ende (M. Schellekens in litt. 2007). One bird wasphotographed at Riung in April 2012 (O. Hidayat in litt. 2013).
The species is recorded regularly by tour groups and individual
birdwatchers at several sites.
Besar: considered locally moderately common (Coates & Bishop
1997). Recorded in semi-deciduous, deciduous and coastal forest(Butchart et al. 1994).
Adonara: two over dry agricultural land and closed forest on
mountain slopes, December 2000 (Trainor 2002); first record forthe island.

Islands are listed west to east.
Lombok: adult pairs observed in Gn Rinjani National Park, andbetween Sembalum and Sapit, in June–July 2003 (J. M. Thiollay inlitt. 2007); the first records for the island. None was reported in areview by Myers & Bishop (2005) or during surveys for the Critically

Pantar: the only record was an adult observed by P. Verbelen at
Gn Wasbila on 3 September 2009 (Trainor et al. 2012).
Alor: observed in Eucalyptus savannah in May 2002 (Trainor
2005a); first record for the island. Some records in 2002, 2009 and102

COLIN R. TRAINOR et al.

2010 by CRT and P. Verbelen may have been confused with Flores
Hawk Eagle, but pairs and singles were seen and confirmed withphotographs at several sites up to about 1,100 m (Trainor et al.
2012).
Timor: single historical specimen, a male (Mees 2006). One
sighting, over mountain forest and peaks at 2,000 m, East Timor,
in 1972 (White & Bruce 1986). Three records in East Timor in1974 (H. Thompson, J. McKean & I. Mason unpubl. data). Since
the 1990s it has been regularly observed in West Timor, particularly
at Bipolo and Camplong (Verbelen 1996, Mauro 1999, Van Biers
2004, N. Kemp in litt. 2007). Noted at four forested localities inWest Timor by Noske & Saleh (1996). CRT had 36 sightings, 25of single birds, 7 of two birds and 4 of three birds, from eight
districts in Timor-Leste over four years 2003–2006; from sea levelto 1,200 m in habitats ranging from coastal flats to village
cultivation, freshwater lakes/swamps, secondary forest and primary
forest (dry deciduous, semi-evergreen, evergreen and montane).
Atauro: pair over montane forest in December 2003, and a captive
juvenile was said to have originated from a nest on Atauro (Trainor& Soares 2004); first records for the island.
Wetar: single historical specimen, originally sexed as male but
probably a female (Mees 2006). Considered locally moderately
common (Coates & Bishop 1997), although this assessment was
based on a half day observation in west Wetar. In 2008, a 44-day
survey recorded it from 8 of 12 sites up to about 500 m: the island
is one of the least disturbed in insular South-East Asia (Trainor etal. 2009); there is at least one additional record of a bird
photographed at sea level in September 2010 (CRT unpubl. data).
Romang: during the first ornithological visit to the island since
1902, a single bird and a displaying pair were observed over tropicalforest at two sites at about 300 m, during two weeks in October2010 (Trainor & Verbelen in press).
Luang: two historical specimens, both males (Mees 2006).
Sermata: one adult bird photographed at forest edge in November
2010 during the first ornithological exploration of the island since
1906 (Trainor & Verbelen in press).
Damar: two sightings over forest and forest edge, August 2006
(Trainor 2007); first records for the island.
Tanimbar islands: first observed on Yamdena, Tanimbar, by
F. Rozendaal between August and November 1985 (F. G.
Rozendaal unpubl. data). Also observed on Yamdena in August
1994 by Verbelen (1996); and in October 1998 by Mauro (1999).
A pair was observed over tall subcoastal primary and secondarysemi-evergreen forest and woodland in January 1996 (Coates &Bishop 1997, Bishop & Brickle 1999). Most recent tour group
records are from the Lorulun area, 20 km north of Saumlaki, butthere are records closer to Saumlaki.
Taxonomic status

In using levels of DNA differentiation to assess taxonomic
assignments of species and subspecies, Norman et al. (1998) andChristidis & Norman (2010) advocated the requirement to include
a relative hierarchical perspective of DNA divergences in the genus
of interest. The relevant DNA data dealing with the taxonomy of
the Aquila fasciata species-complex are summarised below.
The taxonomy of Bonelli’s Eagle and the African Hawk Eagle
Aquila spilogaster has been a contentious issue. Long regarded as asingle species (Brown & Amadon 1968), the recent tendency is to

Forktail 29 (2013)

treat the two as separate species (Thiollay 1994, Ferguson-Lees &Christie 2001). The consistent morphological, plumage andbehavioural differences have been cited as evidence for species-levelseparation. Lerner & Mindell (2005) give a molecular perspective
through their examination of mitochondrial DNA differentiation
in a range of birds of prey, including Bonelli’s Eagle and African
Hawk Eagle. Between the two species there were 16 and 18 basepair differences in cytochrome b and ND2, respectively. Thesefigures were larger than those recorded between other species-pairs
identified in the study: Wedge-tailed Eagle Aquila audax andGurney’s Eagle A. gurneyi; and Little Eagle Hieraaetus morphnoides
and Booted Eagle H. pennatus. Although not conclusive, the DNA
data support separate species treatment for Aquila fasciata and A.
spilogaster.
Apart from the three individuals of A. fasciata examined by Lerner& Mindell (2005), cytochrome b data are available for a further six
individuals. Haring et al. (2007) lodged a 264-base-pair fragment on
GenBank (accession numbers EF459628–EF459631) from two
individuals of A. f. fasciata (one from Italy and one with no locality
information), and two individuals of A. f. renschi from Flores. Helbig
et al. (2005) examined a 1,143-base-pair fragment from an individualof A. f. fasciata from Israel, and Bunce et al. (2005) examined 1,017
base pairs in another A. f. fasciata individual (no locality data). In
addition, JAN & LC sequenced 409 base pairs of an individual A. f.
renschi from Timor, feathers of which were collected by JO. A 217base-pair fragment was common to all five studies, and this wascompared across the three individuals of A. f. renschi and sevenindividuals of A. f. fasciata examined. There were only three variable
sites, and this variation was limited to a unique base change in each of
three individuals of A. f. fasciata. By excluding the four samples from
Haring et al. (2007), it was possible to compare a 267 base-pairfragment across the remaining six individuals, but this did not reveal
any additional variation. There were no differences recorded between
A. f. renschi and the common A. f. fasciata haplotype. The negligible
cytochrome b variation recorded was, therefore, limited tocomparisons within A. f. fasciata. This lack of any moleculardifferentiation between the subspecies A. f. fasciata and A. f. renschi isconsistent with a very recent separation.
Variation in a 253-base-pair fragment of the mitochondrial
control region was assessed in 72 individuals of A. f. fasciata from
Spain, Portugal and Morocco by Cadahía et al. (2007). They found
four mitochondrial types each differing from the other by a single
base-pair change. Moreover, there did not appear to be any
geographic structure to the genetic variation observed across the
populations surveyed. One explanation offered for the low levels ofgenetic variation was a loss of genetic variation caused by population
reduction during the Pleistocene glaciations, and more recently
through human activities such as habitat clearance and hunting.
Control-region sequence data for A. fasciata have also been
lodged on GenBank (Accession Numbers EF459585–459588) by
Haring et al. (2007). Unfortunately, the 237-base-pair fragment
does not correspond to the region examined by Cadahía et al. (2007).
The control-region data of Haring et al. (2007) was obtained fromthe same specimens that cytochrome b data were obtained (seeabove): two individuals of A. f. fasciata (one from Italy and one withno locality information) and two individuals of A. f. renschi from
Flores. The A. f. fasciata individual from Italy differed by 5–6
changes from the other three individuals. The remaining three
individuals differed from each other by 1–2 changes. Although thelevel of variation is low, there is nevertheless more variation recorded
within A. f. fasciata than between A. f. fasciata and A. f. renschi.
Both the cytochrome b and control-region DNA datasets
showed similar patterns: low levels of DNA variation across therange of A. fasciata; no diagnostic DNA marker distinguishing A.
f. fasciata from A. f. renschi; and more variation within A. f. fasciata
than between the two subspecies. Although this pattern of variation

Bonelli’s Eagle Aquila fasciata renschi in the Lesser Sundas, Wallacea

103

COLIN R. TRAINORMARK SCHELLEKENSForktail 29 (2013)

Plate 1. Captive juvenile Bonelli’s Eagle A. f. renschi on Flores, Indonesia,
14 October 2004. The captive bird was kept tethered to a wooden
plank, by a short rope tied to one ankle; the bird had a small woundon the underside of its carpal joint, and an overgrown bill.

Plate 2. Captive juvenile Bonelli’s Eagle in Timor-Leste, 10 April 2003.
Plate 3. Adult Bonelli’s Eagle in flight near Mt Ranaka, Flores, holding
a village chicken, 30 August 2011.

could be indicative of a slow rate of mitochondrial evolution in
Aquila, the comprehensive cytochrome b and ND2 datasets of
Lerner & Mindell (2005) do not provide any such indication. The
widely disjunct distributions of A. f. fasciata and A. f. renschi also
make it unlikely that a lack of differentiation is caused by pastbottlenecks. It is difficult to envisage similar genetic bottlenecks
occurring in such widely separated populations.
Juvenile morphology

Colour photographs (Plates 1 & 2) and other unpublished imagesof the same birds show that juveniles of A. f. renschi are similar inplumage to juveniles of the nominate subspecies, with few evident
differences (V. Hernández in litt. 2007). The photographic
evidence shows colour variation within the range of that of juvenile
Eurasian Bonelli’s Eagles. However, Wallacean birds are more
lightly built than Eurasian birds. Juveniles of each subspecies would
be indistinguishable, and only separable by measurement (V.
Hernández in litt. 2007).
There is little information on the feeding ecology or breeding
biology of Bonelli’s Eagle in Wallacea. A bird was observed feeding
on a Green Junglefowl Gallus varius carcass at Gn Ranaka, Flores,
in August 2007 (Myers 2007), an adult bird was photographedholding a village chicken Gallus sp. near Gn Ranaka in 2011 (Plate3), and in September 2011 an adult Bonelli’s Eagle at Pagal, Flores,delivered a chicken Gallus sp. or a rallid to a juvenile (Robson 2011).
In Ruteng, west Flores, Bonelli’s Eagle were twice (in separate years)
observed flying low over the town, and were suspected to be
searching for village chickens (J. Eaton in litt. 2013). Other likely
prey within their range includes Pink-headed Imperial Pigeon

MARC THIBAULT

Biology

104

COLIN R. TRAINOR et al.

Ducula rosacea, Green Imperial Pigeon D. aenea, Timor Black
Pigeon Turacoena modesta and other forest pigeons, cuscus
Phalanger sp., rats (Muridae) and medium-sized fruit-bats(Pteropodidae) that roost in caves, forests and savannah palms. A
nest was found on 12 July 2003 on Gn Ranaka, Flores, at 1,420 m:
an adult was sitting on the nest in a tree and the mate flew in withprey (J. M. Thiollay in litt. 2013). A second nest was found at
Lermatang on Yamdena, Tanimbar islands, on 22 May 2008 (Yong& Lee 2008): it was about 20–25 m up in a forest tree,
approximately 1.5 m in diameter and consisted of sticks and vines.
It was unclear whether the pair were sitting on eggs or had young,but they were actively managing the nest.

DISCUSSION
This review confirms that Bonelli’s Eagle is more widespread in
the Lesser Sundas than previously believed; recent new records fromnine islands in addition to the nine where it was previously recorded
have extended its range to a land area of about 87,400 km2. The
distinctiveness of the isolated Lesser Sunda population has been
subject to ongoing speculation, due to its smaller size and plumage
differences compared with fasciata; the tail being more strikingly
barred and belly, thighs and crissum more boldly marked(Ferguson-Lees & Christie 2001). However, the negligible levelsof genetic differentiation between A. f. renschi and A. f. fasciata do
not support the contention of Thiollay (1994) and Ferguson-Lees& Christie (2001) that renschi should be accorded full species
status, although it is harder to argue against subspecific recognition
for both forms. The patterns of DNA variation are more consistent
with the relatively recent arrival of renschi in the Lesser Sundas
and it may have been introduced from Eurasia (see below).
Accordingly, the smaller size of renschi and the plumage differences
between it and fasciata would also have evolved relatively rapidly.
Within the archipelago Flores and Timor appear to bestrongholds, with many records from human-modified landscapes,
but this may be partly a result of greater observer effort on theseislands compared with elsewhere. The lack of earlier records from
Sumba and Sumbawa, and recent records from other islands, may
partly reflect bias of historical collectors and recent increase insurvey effort (M. Bruce in litt. 2007). For example, there was areluctance to collect cumbersome large specimens including raptors
because of the relatively high shipping costs (Hartert 1904). Sumba
is relatively well surveyed, and tour parties now visit annually, so
the paucity of records suggests that it is either a rare resident, orthat birds are occasional visitors from nearby islands. Knowledgeof the avifauna of Roti is improving (Verheijen 1975, Trainor
2005b, Collaerts et al. 2011, P. Verbelen in litt. 2010), but there
are no records of Bonelli’s Eagle from this largely deforested island.
Lack of records from other largely deforested islands (Sawu [Sabu],
Semau and Kisar) suggests that a minimum level of forest cover isnecessary to sustain populations of the species. This could be
associated with the scarcity of large prey species, such as frugivorous
pigeons (Newbold et al. 2013), in agricultural land or savannah
woodland.
There are insufficient data to comment on population trends,
but Bonelli’s Eagle appears to be holding its own at present,
although the extensive and rapid deforestation in Indonesia(Brooks et al. 1999) may adversely affect it. The species’s
Indonesian (at the time including East Timor) conservation status
was assessed as ‘no immediate danger’ (van Balen 1994), and there
seems no reason to amend this at present. The species currently
occurs in cultivated lands and secondary forest as well as natural
habitats.
Hunting either for food, or to reduce the perceived impact onvillage livestock, might also affect populations of Bonelli’s Eagle.

Forktail 29 (2013)

Hunting is part of life for many villagers in the Lesser Sundas. On
some islands, hunters are often armed with powerful air-rifles(comparable with a .22 rifle) and children have powerful slingshots.
They shoot a wide range of wildlife, including raptors, and climb
trees to collect nestlings for food. As a predator of village chickens,
Bonelli’s Eagle is likely to be targeted to reduce the perceived impact
on economically important village livestock. Raptor nestlings arecommonly taken captive, and suffer casual, habitual cruelty incaptivity. A captive juvenile Bonelli’s Eagle was photographed
tethered by a metal ring on one ankle attached to a short rope and,not surprisingly, the bird had bumblefoot (a bacterial infection andinflammatory reaction) in the shackled foot as well as cere damage
and abraded carpals (Plate 1). Another captive juvenile/subadultbird owned by a foreign defence worker in Timor-Leste was housed
in a chicken wire cage at a United Nations military compound formonths. It had serious cere damage (CRT unpubl. data). CRT has
also seen at least two other captive Bonelli’s Eagles in Timor-Leste,
although there may be many more out of sight.
Conservation priorities include further field surveys on thelarge islands of Sumba, Sumbawa, Wetar and Yamdena and on
islands where there are no records (e.g. Babar, Moa, Roti, Solorand Lembata). The single largest tropical forest in the Lesser Sundas
is in west Sumbawa (about 2,000 km2) (Jepson et al. 2001), whilst
Wetar retains more than 97% forest cover; both deserve specificsurveys for Bonelli’s Eagle. The breeding biology of this subspecies
is essentially unknown, so it would be useful to monitor populationlevels and breeding success at selected sites on various islands. Thereis also a need for an environmental education campaign todiscourage persecution of eagles in general, improve the lot of
captive birds, and encourage local people to ‘own’ and value suchiconic species (Burnham et al. 1994, Salvador 1994). In TimorLeste, for example, the campaigns should also target foreign
nationals (military and embassy staff). Effective conservation of
Bonelli’s Eagle and other raptors in Wallacea is likely to deliver
broader biodiversity benefits (Sergio et al. 2006).
Origins of the Wallacean population

The isolated populations of Bonelli’s Eagle, and Short-toed SnakeEagle Circaetus gallicus, in Wallacea stand out as zoogeographicallyanomalous—usually explained as relicts from past climatic and sealevel changes (Voris 2000). But it may be important to consider
the human history of these islands; the first Dutch ships arrived inIndonesia (East Indies) in 1596 and determined exploitation
started around 1830. The first Europeans to visit Timor were
Portuguese, perhaps as early as 1512, and the Dutch occupied
Kupang in present-day West Timor in the mid-seventeenth
century, beginning a long conflict for control of the sandalwood
trade. The Dutch controlled most of the Lesser Sundas from the
1600s, but the Portuguese held Flores (especially east Flores with
forts on nearby Adonara and Solor) and East Timor, including theAmbeno (Oecusse) enclave, for long periods (Fox 2003).
Before assuming that the eagle occurs naturally on these islands,
it is important (but difficult) to rule out the possibility that
Bonelli’s Eagles were transported from Europe or South Asia
(notably India where the Portuguese also had colonies) by Dutch
or Portuguese traders or settlers. Europeans may have introduced
eagles to their Indonesian colonies as mascots, pets or falconry birds,
perhaps from Iberia, North Africa or South Asia. Other birds,
junglefowl Gallus spp., and Red Avadavat Amandava amandava
are thought to have been introduced to the Lesser Sundas several
centuries ago. The Red Avadavat is represented in the Lesser Sundas
by the subspecies flavidiventris, which occurs naturally in SouthYunnan (China), Thailand and Myanmar (White & Bruce 1986).
There was also much movement of various animals, by Asian andMelanesian peoples, between Asia, Wallacea and New Guinea(Heinsohn 2003). Alternative hypotheses that need to beForktail 29 (2013)

Bonelli’s Eagle Aquila fasciata renschi in the Lesser Sundas, Wallacea

investigated include vagrant Bonelli’s Eagles from Asia settling in
the Lesser Sundas.

ACKNOWLEDGEMENTS
We thank Vladimir Dinets, James Eaton, Raf Drijvers, Hank Hendriks,
Oki Hidayat, Rob Hutchinson, Nev Kemp, Frank Rozendaal, Mark Schellekens,
Greg Smith, Brian Sykes, Marc Thibault, Jean-Marc Thiollay and Philippe
Verbelen for their unpublished sightings and data, Rui Pires for assisting JO in
the field, and Víctor J. Hernández and James Eaton for evaluating images ofeagles. Thanks go to Will Duckworth and Phil Round for access to an
unpublished manuscript on the South-East Asian population of Bonelli’s Eagle,
and more recent observations. We also thank Mark Schellekens and Marc
Thibault for permission to use their images, and Murray Bruce for commenting
on a draft. SD thanks Sofia Debus for facilitating the initial drafting of thispaper. JO thanks Luis Palma and Marcos Moleón for helpful discussions andfor showing him Bonelli’s Eagles in Iberia.

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Colin R. TRAINOR, Research Institute for the Environment andLivelihoods, Charles Darwin University, Northern Territory 0909,
Australia. Email: halmahera@hotmail.com
Stephen J. S. DEBUS, Division of Zoology, University of New
England, Armidale, NSW 2351, Australia. Email:sdebus@une.edu.au
Jerry OLSEN, Institute for Applied Ecology, University of Canberra,
ACT 2601, Australia. Email: jerry.olsen@canberra.edu.au
Janette A. NORMAN, Department of Genetics, University of
Melbourne, Parkville, Victoria, 3001, Australia. Email:janorman@unimelb.edu.au
Les CHRISTIDIS, National Marine Science Centre, Southern
Cross University, Coffs Harbour, NSW 2450, Australia. Email:les.christidis@scu.edu.au