Landscape characteristics derived from

satellite-tracking data of wintering habitats

used by oriental honey buzzards in Borneo

Syartinilia, Afra D.N.Makalew, Yeni

A.Mulyani & Hiroyoshi Higuchi

Landscape and Ecological

Engineering

ISSN 1860-1871

Landscape Ecol Eng

DOI 10.1007/s11355-013-0237-4

1 23
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 Author's personal copy
Landscape Ecol Eng
DOI 10.1007/s11355-013-0237-4
ORIGINAL PAPER
Landscape characteristics derived from satellite-tracking data
of wintering habitats used by oriental honey buzzards in Borneo
Syartinilia Afra D. N. Makalew Yeni A. Mulyani
Hiroyoshi Higuchi
Received: 18 February 2013 / Revised: 24 September 2013 / Accepted: 23 October 2013
International Consortium of Landscape and Ecological Engineering and Springer Japan 2013
Abstract Knowledge of the spatial distribution of the
wintering habitats of migratory raptors is a prerequisite to
understanding their wintering ecology and managing their
habitats. Oriental honey buzzards (OHBs, Pernis
ptilorhynchus) are migratory raptors with wintering
grounds in Indonesia. OHBs wintering habitats can be
divided into core and edge habitats with unique landscape
characteristics, which influence wintering-site selection.
Twenty-three satellite-tracked OHBs (20062010) used
Borneo as their wintering grounds. The primary aim of this
study was to analyze OHBs wintering habitat distributions
(core and edge habitats) in Borneo and their landscape
characteristics in the province of South Kalimantan. Fixed-
kernel density estimation was used to estimate the edge and
core habitats of 23 OHBs in Borneo. We used a 95 % fixed
kernel (FK_95 %) and a 50 % fixed kernel (FK_50 %) to
estimate the spatial distribution of edge and core habitats,
respectively. Factor analysis was used to analyze landscape
Syartinilia (&)  A. D. N. Makalew
Department of Landscape Architecture, Faculty of Agriculture,
Bogor Agricultural University (IPB), Jl. Meranti, Kampus IPB,
Darmaga, Bogor 16680, Indonesia
e-mail: syartinilia@ipb.ac.id
A. D. N. Makalew
e-mail: amakalew@yahoo.com
Y. A. Mulyani
Department of Forest Resources Conservation and Ecotourism,
Faculty of Forestry, Bogor Agricultural University (IPB), Jl.
Meranti, Kampus IPB, Darmaga, Bogor 16680, Indonesia
e-mail: yenimulyani@ipb.ac.id
H. Higuchi
Graduate School of Media and Governance, Keio University,
Endo 5322, Fujisawa, Kanagawa 252-0882, Japan
e-mail: hhiguchi@sfc.keio.ac.jp
characteristics of core and edge habitats. Results showed
that edge and core habitats covered about 153,463.4 km2
(20.7 %) and 27,528.3 km2 (3.7 %) of the Borneo area,
respectively. Habitat selection by OHBs at wintering sites
in both core and edge habitats was highly influenced by the
availability of thermal winds and food. However, the more
frequent presence of OHBs in core habitats indicates that
habitat selection is basically influenced by increased ther-
mal winds associated with particular landform character-
istics. Identification of these landscape characteristics
provides useful baseline information for ecological-based
development, particularly for landscape management and
biodiversity conservation.
Keywords Core habitat  Edge habitat  Migration 
Pernis ptilorhynchus  Wintering habitat
Introduction
Raptors are at the top of the food chain, so their health
depends on the health of the ecosystems they live in or
migrate through (Meyburg 1986). Thus, declines in raptor
populations can indicate declines in the health of the eco-
systems they depend on. This means that efforts to con-
serve raptors not only ensures that these magnificent
animals continue to live into the future but can also help
preserve entire ecosystems, upon which many other spe-
cies, including humans, depend. Migratory raptors need to
have at least three different environments: breeding
grounds, wintering grounds, and stopover grounds (Bild-
stein 2006).
Indonesia is an important wintering area for several
species of migratory raptors that breed in the Eastern
Palearctic (Germi 2005). However, serious habitat
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destruction, forest fragmentation, and illegal hunting threaten
the survival of important raptors in Indonesia (Syartinilia
2008). The oriental honey buzzard (OHB), which breeds in
southern Siberia, northern Mongolia, northeastern China,
Korea, and Japan, migrates south during the winter (Orni-
thological Society of Japan 2000). The migratory raptor has
winter habitats in Southeast Asia and occurs in the Philip-
pines, Malaysia, Indonesia, and Timor Leste. It is known that
the breeding distribution of this species has contracted and
that its Japanese population is in decline (Higuchi 2005). It is
likely that the same is true in other areas of East Asia;
however, there is very little information available on OHB
populations or on the potential causes of their decline.
Satellite tracking is a powerful research technique that
can be used to study bird movements from mid (within a
country) to large (global) scales. It can provide near real-
time information on the movements of individual birds
anywhere in the world (Cohn 1999; Webster et al. 2002;
Higuchi and Pierre 2005). This tracking can provide
information on the precise location of migration routes, the
temporal schedule of migration, and the relative impor-
tance of stopover sites based on the number of bird visits
and the duration of their stays. In Asia, satellite-tracking
studies have been carried out intensively since the 1990s,
and much information on migration of cranes, storks,
waterfowl, and raptors has been gathered (e.g., Higuchi
et al. 1994, 1996, 1998, 2004, 2005; Kanai et al. 1997,
2000, 2002). Satellite tracking has been successfully used
to track the OHB since 2003. Satellite-tracking methods
radically extend detection ranges compared with conven-
tional techniques used to track migrant species that
undertake intercontinental and even transoceanic journeys.
This technology is increasingly used in raptor-migration
science (Bildstein 2006). In general, satellite tracking alone
only provides time and location data (when and where an
animal appears). However, by integrating this data with
environmental variables and coupling with geographic
information system (GIS) techniques, it is possible to
analyze the habitat use of tagged birds. Researchers have
used this approach to examine migration strategies (Fujita
et al. 2004), study habitat use (Kernohan et al. 1998), and
analyze interspecific competition for food resources (Bar-
low et al. 2002). These studies have revealed OHB
migration routes (Higuchi et al. 2005), site fidelity (Shiu
et al. 2006), and shifts in migratory routes (Yamaguchi
et al. 2008). However, few studies have focused on the
spatial distribution and landscape characteristics of win-
tering, stopover, and breeding habitats used by OHBs. This
information is vital for developing effective conservation
strategies for OHBs.
A total of 49 OHBs have been tracked by satellite since
2003, and about 47 % (23 OHBs) of the individuals tracked
have used Borneo as their wintering site (Higuchi et al., in
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preparation). Therefore, Borneo is an ideal location in
which to study the wintering habitat characteristics of
OHBs. Borneo has the highest percentage of OHB win-
tering habitats, and thus, its preservation is critical for the
conservation of this species. However, Borneo has expe-
rienced heavy deforestation and forest degradation during
the past two decades. Transmigration to the Indonesian part
of Borneo and poorly planned development projects such
as the Mega Rice Project (MRP) have strongly contributed
to the loss of natural forest ecosystems and resulted in
increased fire risk associated with shifting cultivation
activities (Jepson et al. 2001; Page et al. 2002; Denis and
Colfer 2006)., Indonesian government sources report that
between 1985 and 1997, Kalimantan lost nearly 10 million
ha of forest, which was the largest absolute loss among
Indonesias major islands [World Resources Institute
(WRI) 2002]. Assuming linear rates of change, the annual
deforestation rate in Kalimantan was 2.1 % over this per-
iod, approximately three times greater than the rate
reported for Southeast Asia as a whole (Archard et al.
2002). A recent study that used a moderate resolution
imaging spectroradiometer (MODIS) to monitor land-cover
change in Borneo between 2002 and 2005 (Langner and
Miettinen 2007) found that in 2002, approximately 57 % of
the land surface of Borneo was covered with forest, of
which 74 % was dipterocarp and [23 % peat-swamp for-
est. The average annual deforestation rate between 2002
and 2005 was 1.7 %, and the carbon-rich peat-swamp
forest ecosystem had a deforestation rate of 2.2 %. Almost
98 % of all deforestation caused by forest fires occurred
within a range of 5 km from the forest edge (Langner and
Miettinen 2007). Although Kalimantans original forest
cover remained higher than on the other Greater Sunda
islands of Sumatra and Java, the high commercial timber
value of Kalimantans lowland dipterocarp forests (DLFs)
has attracted significant investment in capital-intensive
logging. This has stimulated the growth of domestic ply-
wood production, accelerated commercial logging, and
expanded plantation agriculture, especially oil palm (Elaeis
guineensis) (Barber and Schweithelm 2000; WRI 2002). It
is likely that the loss of forests and habitat deterioration
will cause population declines in migratory raptors such as
OHBs.
Understanding landscape habitat characteristics of win-
tering OHBs is a prerequisite to understanding their win-
tering ecology and managing that habitat. OHBs have both
core and edge habitats, which are considered to have their
own landscape characteristics that influence the way that
OHBs use them as wintering sites. The definition of a core
habitat varies depending on species and habitat require-
ments (Semlitsc and Jensen 2001; Porej et al. 2004; Goetz
et al. 2009). Generally, a core habitat has a high degree of
isolation and is surrounded by edge habitat that acts as a
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buffer area. The buffer area plays an important role as a
corridor for maintaining the quality of its core habitat
(Phillips 2002). In this study, we define core habitat as an
area with a high proportion of occupation based upon
satellite tracking data sets. A major problem facing con-
servation efforts is to accurately identify core habitats of
species. This is an important part of habitat management
and can enable protection and restoration of habitats affected
by anthropogenic activities. Using appropriate techniques,
fixed-kernel density function coupled with GIS can be an
effective first step in identifying core habitats.
The aims of this study were to: (1) produce a spatial
distribution map of core and edge wintering habitats of
OHBs in Borneo, (2) determine the size of core and edge
wintering habitats, and (3) identify landscape characteris-
tics of both core and edge habitats in the province of South
Kalimantan. Results of this study will provide valuable
information on the spatial distribution (location and size)
and landscape characteristics of OHB wintering habitats,
which applies to the conservation and management of their
wintering habitats. Finally, the method used in this study
can also be applied to other species.
Fig. 1 Study area
Materials and methods
Study areas
The study was conducted on Borneo Island, the third largest
island in the world (Fig. 1). The island is divided between
three countries: Brunei, Indonesia, and Malaysia, with a
total area of 746,305 km2. Kalimantan is the name given to
the Indonesian part of Borneo, which covers 73 %
(544,802 km2) of Borneos land mass. Kalimantan has four
provinces: West Kalimantan, Central Kalimantan, South
Kalimantan, and East Kalimantan. More than half the island
lies below 150 m in altitude and has no active volcanoes;
however, its main mountain ranges are igneous in origin.
Kalimantan lies on the equator in a region that experiences
high temperatures throughout the year and within the wettest
part of the Indonesian Archipelago. These conditions and the
islands geological and climatic history have encouraged
speciation and high species diversity. Borneo has at least
3,000 species of trees, including 267 species of dipterocarps
(58 % are endemics), the most important group of commer-
cial timber trees in Southeast Asia (Mackinnon et al. 1996).
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A case study of the landscape characteristics of OHB
wintering habitats was conducted in the province of South
Kalimantan, Indonesia (Fig. 1), a province 38,744.23 km2
situated between latitude S1210 S4100 and longitude
E114190 E116330 . The provincial capital is Banjarma-
sin, and its boundaries are Makassar Strait in the east,
Central Kalimantan in the west and north, the Java Sea in
the south, and a small part of East Kalimantan in the north
(Mackinnon et al. 1996). Topography is mostly below
300 m asl (88.06 %), with slopes below 3 % (39.78 %)
based on Advanced Spaceborne Thermal Emission and
Reflection Radiometer Global Digital Elevation Model
(ASTER GDEM) data. Analysis of images captured by
Advanced Land Observing Satellite Phased Array type
L-band Synthetic Aperture Radar (ALOS PALSAR)
showed that lowland DLF and agriculture/plantation
covered about 35.94 and 19.44 % of this province,
respectively, in 2007. The population of South Kaliman-
tan was recorded at just over 3.626 million people in
2010, with about 41.66 % working in the agriculture
sector and a population growth rate over the last decade of
1.98 % [Statistic Indonesia (BPS) 2010]. Fieldwork was
conducted in February 2011. Two core habitats (Sultan
Adam and Hulu Sungai Selatan) and three edge habitats
(Sultan Adam, Hulu Sungai Selatan, and Barito River)
were also visited at that time.
Data
Wintering sites
We located the honey buzzards through the Argos system
(Argos 1996). Argos assigns a location class (LC) to each
of the estimated locations based on their accuracies.
These are then ranked from Z (least accurate), B, A, 0, 1,
2, to 3 (most accurate). Accuracies for the least accurate
LCs, A, B, and Z are filtered out by the Argos system. In
general, we used LC 03 for data analysis, which have
errors of about 1,000 m, 3501,000 m, 150350 m, and
\150 m, respectively. LC A and B data were included
only when they were spatially and temporally close to LC
03 category based on the time and distance moved.
Wintering sites were defined following Higuchi et al.
(2005) as the area \30 km in diameter for at least 24 h.
We used satellite-tracking data from 23 adult OHBs,
consisting of 11 females and 12 males, from three dif-
ferent breeding sites in Japan, which have been tracked
from 2006 to 2010. Among the 23 OHBs, five birds wore
transmitters with a GPS unit; four of them had their
wintering areas in South Kalimantan Province. In this
study, 3,193 wintering sites throughout Borneo were
identified and analyzed using the information from 23
OHBs (Fig. 2).
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Environmental variables
The selection of environmental variables is based on the
basic need of the migratory raptors in their wintering
grounds: food availability (Hutto 1985) and thermal wind
(Bruderer et al. 1994). All environmental variables were
derived from elevation, slope, and land-cover maps, which
were converted into Euclidean distance images. To produce
Euclidean distance images, we first produced a binary map
of each elevation class, slope class, and land-cover type
then converted them to vector maps and finally calculated
the Euclidean distance. We calculated 21 environmental
variables composed of terrain factors (five and six
Euclidean distance images to the nearest border of eleva-
tion and slope classes, respectively) and ten Euclidean
distance images to the nearest border of neighboring land-
cover patches (Table 1). All image layers were prepared
and analyzed as raster data with a pixel size of 50 9 50 m
using ArcGIS 9.3.1 and ERDAS Imagine 9.1.
Spatial distribution analysis
One of the most widely used methods for spatial distribu-
tion analysis is kernel-density estimation (KDE). KDE
creates contours of utilization intensity (isopleths) by cal-
culating the mean influence of data points at grid inter-
sections (Worton 1989) and is based on the assumption that
the aggregation of locations provides information about the
differential use of space within the home range of a species
(Anderson 1982; Worton 1989; Kernohan et al. 2001).
Each isopleth contains a fixed percentage of the utilization
density that corresponds to the amount of time the animal
spends in the area enclosed by that contour (Hemson et al.
2005). This is based on the assumption that an animal
uniformly uses the area inside a contour enclosing a certain
proportion (e.g., 95 %, 50 %) of the total probability
density of the recording locations derived from the satel-
lite-tracking data.
The fixed-kernel density technique was selected because
it is considered the most robust of the various home-range
estimators (Worton 1989; Seaman and Powell 1996;
Wauters et al. 2007) and has frequently been used in
habitat selection studies (Mace et al. 1996; Mace and
Waller 1997; Kenward et al. 2001; McLoughlin et al. 2002;
Lyons et al. 2003). One of the aims in this study was to
define both core and edge habitats, where the core is
defined as an area that has a higher proportion of occupa-
tion than edge habitats. Even though we did not estimate
home-range sizes, we used the kernel method to delineate
core and edge wintering habitats used by OHBs based on
satellite-tracking data. We used a 95 % fixed kernel
(FK_95 %) to estimate the spatial distribution of edge
habitats and a 50 % fixed kernel (FK_50 %) to estimate the
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Fig. 2 Satellite-tracking data
showing the location of
wintering sites of the 23 oriental
honey buzzards (OHBs) tracked
from 2006 to 2010
distribution of core habitats of the 23 OHBs in Borneo.
This analysis was performed using the Hawths tools
(http://www.spatialecology.com) as a free extension for
spatial ecology tool for animals in ArcGIS 9.3.1. Then, we
overlaid the obtained spatial distribution of core and edge
habitats with the existing protected area boundaries (World
Database on Protected Areas, http://www.wdpa.org/
Download.aspx) in order to determine the legal status of
that habitat.
Analysis of landscape characteristics
Habitat is defined as conditions and resources present in an
area that produce occupancy (including survival and
reproduction) by an organism (Hall et al. 1997). In essence,
habitat is a multivariate concept (Morrison et al. 1992;
Youlatos 2004; Praca et al. 2009), and many statistical
methods have been developed to model habitat selection,
such as modeling the probability that an organism will use
a resource unit (e.g., a pixel of a raster map) based on the
environmental variables measured for that unit. These
methods are widely used, as they rely on the generalized
linear model, which is available in most standard statistical
packages. Most of these modeling methods require prior
knowledge of the landscape characteristics that can influ-
ence the distribution and abundance of the species being
studied (Boyce and McDonald 1999). One of the most
difficult tasks is to decide which explanatory variables or
combination of variables (Guisan and Zimmermann 2000)
is influencing habitat selection. The modeling step should
be preceded by an exploration step (Calenge et al. 2005;
Calenge 2007) using statistical methods such as factor
analysis.
Factor analysis was used to integrate 21 environmental
attributes and identify landscape characteristics of core and
edge habitats used by OHBs. Factor analysis has frequently
been used in habitat selection research, including studies on
African elephants (de Knegt et al. 2011), red-back shrike
(Titeux et al. 2007), and houbara bustards (Osborne et al.
1997). Prior to carrying out factor analysis, two-sample
t tests (P \ 0.05) were performed on environmental vari-
ables in each core and edge habitat; A probability of B0.05
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Table 1 Environmental
No. Variables Variable Source
variables used in this study
abbreviation

1. Nearest distance to elevation 0300 m DE1 ASTER Global Digital Elevation

2. Nearest distance to elevation 300500 m DE2 Model (ASTER GDEM)
3. Nearest distance to elevation 500700 m DE3 Available at: http://www.gdem.
aster.ersdac.or.jp/
4. Nearest distance to elevation 7001,000 m DE4
Resolution: 30 9 30 m
5. Nearest distance to elevation [1,000 m DE5
6. Nearest distance to slope 03 % DS1 Generated from ASTER GDEM
7. Nearest distance to slope 38 % DS2
8. Nearest distance to slope 815 % DS3
9. Nearest distance to slope 1525 % DS4
10. Nearest distance to slope 2540 % DS5
11. Nearest distance to slope [40 % DS6
12. Nearest distance to build-up area DBA Land-cover map
13. Nearest distance to bare land DBL Source: Laboratorium of Remote
14. Nearest distance to oil palm forest DOF Sensing and GIS, Faculty of
Forestry, Bogor Agricultural
15. Nearest distance to paddy field DPF
University
16. Nearest distance to swamp bush DSB
Image source: ALOS PALSAR,
17. Nearest distance to agriculture/plantation/shrubs DAP 2007
18. Nearest distance to mangrove forest DMF Resolution: 50 m 9 50 m
19. Nearest distance to water body DWB
20. Nearest distance to dry land forest DLF
21. Nearest distance to peat swamp forest DPS

was accepted as being significant, and variables were
retained for further analysis. The preliminary t test is
intended to reject the null hypothesis that there are no
significant differences in characteristics of core and edge
habitats. Subsequently, factor analysis was carried out with
Varimax rotation with the Kaiser normalization to extract
factors that characterize core and edge habitats. To extract
the values of environmental variables from each pixel in
the Euclidean distance images, we created sampling points.
Sampling points were selected randomly in both core and
edge habitats using the Hawths tool extension in ArcGIS
9.3.1. About 27,434 and 151,392 sampling points were
taken from core and edge habitats, respectively. Factor
analysis was carried out separately on environmental
variables values extracted from core and edge habitats
using SPSS 15 (SPSS Inc.)
Results
Spatial distribution and size of wintering habitat used
Spatial distribution of habitats used by OHBs are shown in
Fig. 3. Using the FK_95 % and FK_50 % for the entire
study period, we obtained a habitat size ranging from 224
to 26,504 km2, 15 to 5,820 km2, with an overall mean
habitat size of 7,481 1,737 (S.E.) km2 and 1,214 329
(S.E.) km2, respectively. Total size of core and edge hab-
itats estimated by FK_95 % and FK_50 % covered about
153,463.4 km2 (20.7 %), and 27,528.3 km2 (3.7 %) of the
Borneo area, respectively. The core area was located
mainly over Sabah, Central Kalimantan, South Kalimantan,
and around the northern boundary of Sarawak and East
Kalimantan. Comparing the core wintering habitat to
existing protected area boundaries showed that only
4,672 km2 (16.7 %) of the core habitat is protected; the
remaining area extends far outside the protected areas. The
edge was distributed around core habitat.
Landscape characteristics
All environmental variables (21 variables) were signifi-
cantly different (P \ 0.05) between core and edge habitats.
The first of five factors explained 79.2 % and 72.8 % of the
observed variation in landscape characteristics of core and
edge habitats, respectively (Tables 2 and 3). Based on
these results, we focused on the first three factors in each
core and edge habitat, which explained about 66.2 % and
59.6 % of data variance, respectively.
This analysis revealed that core habitats were charac-
terized by a number of landscape characteristics. The first
factor indicated that core landscapes were characterized by
having an elevation [300 m (DE2DE5), a slope [15 %
(DS4DS6), and influenced by the nearest distance to

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Fig. 3 Spatial distribution map
of oriental honey buzzard
(OHB) core and edge wintering
habitats in Borneo from 2006 to
2010 based on fixed-kernel
methods and overlaid with
protected area boundaries
lowland DLF. The second factor showed that core habitats
were also associated with particular land-cover types, such
as swamp bush (DSB) and paddy fields (DPF), which is
likely to be related to the distribution of OHB food
resources. The third factor was related to flat to steeper
slopes (DS1DS3). Analysis of edge habitats revealed the
first factor was related to landform and particularly to
elevations [300 m (DE2DE5) and main foraging area
(DLF). Unlike core habitats, slope was not included in the
first factor for edge habitats. The second factor was mainly
related to the distribution of food resources and was
characterized by various land-cover types, including
swamp bush (DSB), paddy fields (DPF), and agriculture/
plantation/shrubs (DAP). The third factor was related to
slopes [15 % (DS4DS6).
Discussion
The characteristics of core and edge OHB wintering hab-
itats were compared in order to increase our understanding
of which factors are most important when developing
wintering habitat management plans for OHBs. A large
amount of variation in habitat selection between core and
edge habitats was explained by the first and second factors.
Landscape characteristics in both habitats were predomi-
nantly influenced by the first factor, which explained about
44.3 % and 35.1 % of data variance, respectively. The first
factor was composed of two environmental variables with
equal importance in both core and edge habitats, i.e., ele-
vation [300 m and nearest distance to the lowland DLF;
and one environmental variable, which differentiated core
and edge habitat, i.e., slope [15 % (Fig. 4). Slope also had
a higher influence in core habitat than in edge habitat, as
slope was not included in the first or second factors for
edge habitat.
Slope and elevation are the physical variables that
comprise the topography of an area. A combination of
landform features and favorable weather generates thermal
winds [Asian Raptor Research & Conservation Network
(ARRCN) 2012]. The presence of thermal winds both in
core and edge habitats can influence bird distributions due
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Table 2 Factor-analysis results for core habitat Table 3 Factor loading of the principal components analysis (PCA)
used for edge habitat
Variables Factor
Variables Factor
1 2 3 4 5
1 2 3 4 5
DBA 0.729
DBL 0.801 DBA 0.723
DOF DBL 0.846
DPF 0.705 DOF
DSB 0.821 DPF 0.793
DAP DSB 0.564
DMF 0.552 0.524 DAP 0.606
DWB 0.768 DMF
DLF 0.634 0.643 DWB
DPS 0.847 DLF 0.625
DS1 0.579 DPS 0.707
DS2 0.960 DS1
DS3 0.960 DS2 0.955
DS4 0.743 DS3 0.955
DS5 0.715 DS4 0.762
DS6 0.523 DS5 0.816
DE1 0.547 DS6 0.825
DE2 0.871 DE1
DE3 0.894 DE2 0.902
DE4 0.873 DE3 0.929
DE5 0.838 DE4 0.934
Eigenvalue 9.311 2,912 1.685 1.505 1.219 DE5 0.928
Variability (%) 44.339 13.869 8.022 7.168 5.804 Eigenvalue 7.372 3.146 1.993 1.634 1.143
Cumulative (%) 44.339 58.208 66.229 73.397 79.202 Variability (%) 35.107 14.980 9.488 7.783 5.445
Cumulative (%) 35.107 50.087 59.575 67.358 72.803
Rotation method: Varimax with Kaiser normalization. Values are
omitted if \0.5. See Table 1 for variable definitions Rotation method: Varimax with Kaiser normalization. Values are
omitted if \0.5. See Table 1 for variable definitions
to their effects on flight behavior. During migration, raptors
move largely by soaring and gliding, using thermal winds et al. 2008). Subsequently, their wintering habitat selection
to save energy (Panuccio 2011). Bildstein (2006) men- will also depend on the habitat preferences of bees and
tioned that land-based thermals are influenced by variation particularly the location of trees with bee-colony nests.
in terrain surfaces and solar radiation. This might explain Starr et al. (1987) reported that Apis dorsata in Borneo
the association we found between slope and core habitats. preferred to build huge honeycombs in the forest on very
Increased slopes in core habitats will provide more thermal tall trees free of epiphytes, or lianas with branches [15 m
winds necessary for soaring OHBs. Bruderer et al. (1994) long. The most common tree occupied is Koompassia sp.,
found that OHBs in southern Israel use thermals in their one of the worlds tallest known angiosperm tree species.
flight, allowing them to save energy for long migrations. The Mate0 -mate0 Dayak people, one of local people living
Wintering habitat selection by migrants might also be in Kalimantan, report that six tree species are inhabited by
influenced by local food abundance and the presence of honeybees, including several dipterocarps (e.g., Shorea
competitors (Petit 2000; Moore and Aborn 2000). The laevifolia), but the most important are K. excelsa and K.
nearest distance to the lowland DLF was included in first malaccensis (De Jong 2000), both of which are widely
factor for both core and edge habitats. This could be related recognized as dominant trees that throughout forests in
to food availability in local forests. According to Hutto Kalimantan. Bees and wasps mostly build nests in the
(1985), food is the major force driving bird habitat selec- forests and feed in surrounding agricultural land or plan-
tion during the nonbreeding season. Therefore, finding a tation areas. The distribution of food resources influences
place with an abundance of food is likely to be one of the habitat selection by OHBs both in core and edge habitats.
main aims of migrating OHBs. OHBs feed on bees and This is supported by the second factor, which was associ-
wasps, such as giant honeybees Apis dorsata (Yamaguchi ated with nearest distance to agriculture land.

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Landscape Ecol Eng
Fig. 4 Typical core (a) and edge (b) habitat in South Kalimantan
Province
Our results suggest that the selection of both core and
edge wintering habitats by OHBs is highly influenced by
the presence of thermal winds and food availability.
However, the more frequent presence of OHBs in core
habitat indicates that habitat selection is particularly
influenced by increased thermal wind availability associ-
ated with certain landform characteristics. These landscape
characteristics can provide useful baseline information for
ecology-based development, especially for landscape
management and OHB conservation. Most of the core
habitat was located outside protected areas (83.3 %), and
this should be taken into account by agencies (i.e., minis-
tries, national parks, nongovernmental organizations) when
developing conservation plans to preserve the wintering
habitat of migratory raptors in Borneo, especially OHBs.
To preserve this habitat, cooperation between stakeholders
is essential, including local people, policymakers, and
landscape architects.
Acknowledgments The authors express sincere thanks to T. Endo,
N. Hijikata, E. Hiraoka, M. Hotta, K. Kuno, H. Nakamura, F. Na-
kayama, Y. Nishizawa, M. Saeki, M. Takahashi, K-I. Tokita, K.
Uchida, and A. Uematsu for their field assistance. We thank Dr.
S. Kitamura and Dr. H. Samejima who gave us information on
tropical forests. We are grateful to the Raptor Indonesia (RAIN),
especially to Zaini Rakhman, Abit, Adam A. Supriatna, for their
guidance during field surveys and also comments on early manuscript
drafts. We give special thanks to our student Annisa Hasanah for her
great help in this study. Finally, we are grateful to Mrs. Diane V.
Wildsmith for proofreading the manuscript. This research was funded
by the Ministry of the Environment in Japan and the Directorate
General of Higher Education (DGHE) in Indonesia under the Inter-
national Research Collaboration and Scientific Publication grant.
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