B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4

available at www.sciencedirect.com

journal homepage: www.elsevier.com/locate/biocon

A gap analysis of Southeast Asian mammals based

on habitat suitability models

Gianluca Catulloa, Monica Masia, Alessandra Falcuccib, Luigi Maioranob,*,

Carlo Rondininib, Luigi Boitanib
a
Istituto di Ecologia Applicata, via Bartolomeo Eustachio 10, 00161 Rome, Italy
b
Department of Animal and Human Biology, Sapienza Università di Roma, Viale dell’Università 32, 00185 Rome, Italy

A R T I C L E I N F O A B S T R A C T

Article history: Southeast Asia is one of the richest reservoirs of biodiversity on earth and home to one of
Received 9 November 2007 the highest concentrations of endemic species. Many protected areas (PA) have been estab-
Received in revised form lished across the region, but to date no systematic evaluation of their efficacy has been
7 August 2008 published because no comprehensive dataset was available which could be fed into an
Accepted 11 August 2008 analysis of conservation gaps. We collected the geographic range for 1086 mammal species
Available online 23 September 2008 of Southeast Asia and we built species-specific habitat suitability models for 901 of them.
We performed two gap analyses (one based on a combination of distribution models and
Keywords: distribution ranges and one based on distribution ranges only) for each mammalian spe-
Southeast Asian mammal databank cies, to evaluate the effectiveness of the existing network of PA and to identify priority
Gap analysis regions and priority species for expanding and consolidating the network. Our results indi-
Protected areas cate that 7.5–8.2% of species are not covered by any PA, and 51.6–59.1% are covered only
Distribution models partially. These species are distributed throughout the entire study area and their conser-
Mammal distribution ranges vation requires the creation of new PA that can help fill this existing conservation gap. This
Threatened species would be particularly important for species which are endemic of small islands, where
species survival is often threatened by the presence of introduced species and habitat con-
version. Yet PAs cannot be considered as the ending point of a conservation strategy,
because overall, 34% of the species we analyzed (many of which already covered by exist-
ing PAs) were at risk of extinction when considering the IUCN red-list criteria. PAs should
therefore be considered in a broader framework of all local ecological and socio-economic
trends, including the growing human population, growing economy and infrastructure
development.
Ó 2008 Elsevier Ltd. All rights reserved.

1 Introduction country system composed by Indonesia, Malaysia and the

Southeast Asia is one of the richest reservoirs of biodiversity
on earth and home to one of the highest concentrations of en-
demic species (Sodhi et al., 2006a). The region encompasses
four hotspots (Myers et al., 2000), several of the most valuable
eco-regions (Olson and Dinerstein, 1998), and a megadiversity
Philippines (McNeely et al., 1990). This extraordinary species
richness encompasses all taxa, and mammals are no excep-
tion (Brooks et al., 1999; Sodhi and Brook, 2006). In fact, roughly
one quarter of the mammal species of the world occurs in the
area, with many new families and species which have been
discovered recently (Jenkins et al., 2005; Musser et al., 2005).

* Corresponding author: Tel.: +39 0649694262; fax: +39 06491135.

E-mail addresses: g.catullo@ieaitaly.org (G. Catullo), m.masi@ieaitaly.org (M. Masi), alessandra.falcucci@uniroma1.it (A. Falcucci),
luigi.maiorano@uniroma1.it (L. Maiorano), carlo.rondinini@uniroma1.it (C. Rondinini), luigi.boitani@uniroma1.it (L. Boitani).
0006-3207/$ - see front matter Ó 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2008.08.019
 B I O L O G I C A L C O N S E RVAT I O N
The rapid and extensive destruction of habitats occurring
worldwide across the tropical belt has not spared Southeast
Asia, which indeed has one of the highest relative rates of
deforestation among the major tropical regions (Laurance,
1999). Its native biota is seriously threatened by forest conver-
sion, forest fires, unsustainable subsistence hunting and
wildlife trade (Sodhi et al., 2004a). In this context, protected
areas (PAs) constitute a valuable tool for assuring the conser-
vation of viable populations within natural ecosystems (Bru-
ner et al., 2001; Redford and Richter, 1999; Groves, 2003;
Rosenzweig, 2003; Cardillo et al., 2006; Lee et al., 2007). Over
the years, Southeast Asian countries have established
(though at different paces) national PA systems. However,
PAs are often not completely representative of the biodiver-
sity of a region (Pressey et al., 1993; Scott et al., 1993; Rodri-
gues et al., 1999; Margules and Pressey, 2000; Rondinini
et al., 2005) as their location and design is frequently based
on socio-economic, aesthetic and political values rather than
biological criteria. Consequently, unrepresentative sites of
lesser conservation value may be set aside for conservation
while sites of the higher value remain unprotected.
Gap analysis (assessing to what extent animal and plant
species are being protected) is a powerful approach to explore
the effectiveness of a PA system in representing local biodi-
versity (Scott et al., 1993). Gap analysis has been applied to
various taxa globally (Rodrigues et al., 2004a), at continental
level, and in many countries worldwide (Scott et al., 1993,
2001; Fearnside and Ferraz, 1995; Ramesh et al., 1997; Rodri-
gues et al., 1999; Powell et al., 2000; De Klerk et al., 2004; Fjeld-
sa et al., 2004; Oldfield et al., 2004; Yip et al., 2004; Dietz and
Czech, 2005; Maiorano et al., 2006, 2007; Rondinini et al.,
2006a), but so far it has never been applied to PAs of Southeast
Asia because no comprehensive dataset has been available
which could be fed into the GAP process.
Rodrigues et al. (2004b) and Brooks et al. (2004) suggested
the necessity of fine scale mapping works that, starting from
the results of the global gap analysis, provide the local context
that is necessary for conservation. The Southeast Asian Mam-
mal Databank project (SAMD – a joint effort between Istituto
di Ecologia Applicata, European Commission and IUCN) for
the first time provides a comprehensive, fine-grained, and
large scale biodiversity dataset, consisting of extent of occur-
rence maps, species–habitat relationships and habitat suit-
ability models for all mammals of the region. The dataset
was compiled in 2002–2006 in close collaboration with the
IUCN Global Mammal Assessment process and it is freely
available at SAMD’s website (http://www.ieaitaly.org/samd/).
In this paper we present the SAMD dataset and provide an
assessment of the effectiveness of the existing PAs for the
conservation of terrestrial mammals in Southeast Asia. While
geographic ranges grossly overestimate species distribution
(conditional on species range size as suggested by Jetz et al.,
2008), thus hiding a number of gap species that overlap PAs
only in unsuitable portions of their range (Rondinini et al.,
2005), distribution models may potentially underestimate
species distribution, providing an overly restrictive picture
of the current conservation status. As suggested by Rodrigues
et al. (2004b) and by Brooks et al. (2004), we focused on the
overlap of PAs with estimated (and validated) suitable areas
for species, because these are more likely to host the species
1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2731
under analysis (Rondinini et al., 2006b). However, we per-
formed our analyses also considering geographic ranges to
obtain a range of reasonable estimates for gap species. Our
analysis is unique in providing indications of conservation
status for a large study area and at a scale that is useful for
conservation planning purposes.
2. Materials and methods
Our study area encompasses the entire Southeast Asia,
including all countries south of China and east of India: Bru-
nei Darussalam, Cambodia, Indonesia, Lao PDR, Malaysia,
Myanmar, Papua New Guinea, The Philippines, Singapore,
Thailand, and Viet Nam.
We characterized the landscape of the study area consider-
ing datasets from different sources and covering: (a) land-cov-
er (Global Land Cover 2000 [GLC2000]; European Commission,
Joint Research Center, 2003; http://www-gvm.jrc.it/glc2000);
(b) elevation (Hastings et al., 1999; http://www.ngdc.noaa.-
gov/mgg/topo/globe.html); (c) hydrology (Digital Chart of the
World; ESRI, 1993); (d) human settlements (Digital Chart of
the World; ESRI, 1993); (e) administrative boundaries (Digital
Chart of the World; ESRI, 1993); (f) PAs (World Database on
Protected Areas [WDPA] Consortium, 2006; http://maps.geo-
g.umd.edu/WDPA/WDPA_info/English/index.html). All layers
were re-sampled for the analyses using a common origin
and a 1-km cell size. From the WDPA, we selected 1635 PAs
presently mapped for SE Asia, irrespective of their IUCN clas-
sification (IUCN, 1994). From analyses, we excluded 90 PAs for
which no information was available on their boundaries and/
or on their area. Of the 1635 PAs, 1088 had information on
their geographical boundaries, while 547 had the coordinates
of their geographical center and the information on the size
of the PA. In order to merge all PAs into a single layer, we built
circular buffers of the same size as the PA around these cen-
tral points.
We compiled a geographical database (freely available
through SAMD’s website) covering the entire study area with
information on 1086 mammal species, belonging to 17 orders
(Table 1).
For each species, we collected from the scientific literature
all available information on the species–habitat relationships,
the elevation range, and the Extent of Occurrence (EOO; the
full list of papers is available from the authors upon request).
Species–habitat relationships (sensu Corsi et al., 2000) fol-
lowed the GLC2000 land-cover classes, and each class was
classified as suitable (main or preferred habitats) or unsuit-
able (secondary and unsuitable habitats). Elevation was clas-
sified according to two classes: inside known elevation
range, outside known elevation range.
All the information collected was reviewed, discussed, and
integrated during five dedicated workshops which involved a
selection of the most eminent local and international experts
on various taxa. Each workshop involved from 21 to 48 ex-
perts for 3–5 days of intensive analysis and updating of the
available data (Mammals of Southeast Asia – Thailand, May
2004; Mammals of the Philippines – Philippines, April 2006;
Bats and Large Mammals of Southeast Asia – Indonesia,
May 2006; Small Carnivores of Southeast Asia – Vietnam, July
2006; Primates of Southeast Asia – Cambodia, September
2732 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4

Table 1 – Orders and number of species included in the Table 2 – Overlay of environmental layers
Southeast Asian Databank
Land-covera Elevationb Suitability score
Order All species Threatened species
1 1 Suitable
Monotremata 4 3 1 2 Not-suitable
Dasyuromorphia 16 3 2 1 Not-suitable
Peramelemorphia 12 2 2 2 Not-suitable
Diprotodontia 58 30
a Suitability scores for land-cover as defined in the method sec-
Proboscidea 1 1
tion: 1 = suitable; 2 = unsuitable.
Scandentia 17 11
b Suitability scores for elevation as defined in the method section:
Dermoptera 2 1
1 = inside known elevation range; 2 = outside known elevation
Primates 76 62
range.
Rodentia 388 114
Lagomorpha 6 1
Erinaceomorpha 7 1 area of potential species presence. The complete species list is
Soricomorpha 56 5 available at SAMD’s website (http://www.ieaitaly.org/samd).
Chiroptera 328 73
Pholidota 3 3
Carnivora 54 24
2.2. Validation
Perissodactyla 3 3
Artiodactyla 52 34 To test the predictive power of the DMs, we measured the per-
Total 1086 371 formance of the models in predicting species potential pres-
Threatened species include critically endangered, endangered, ence. In particular, we measured the agreement between
vulnerable, and near threatened. each DM and a set of points of presence which were indepen-
dently collected in the field. We also measured the agreement
between each DM and a set of random points and we com-
pared the results using a permutation test.
2006). Our dataset was also integrated with the information We obtained point data on species presence gathering the
gathered during two additional workshops run by IUCN in available published and unpublished datasets, consisting
the framework of the Global Mammal Assessment project mainly in observations and captures. In particular, we ob-
(Mammals of Australia and Papua New Guinea – Australia, tained data for Chiroptera, Rodentia, Artiodactyla, Carnivora,
August 2005; Rodents of Southeast Asia - United States, May Primates and Scandentia covering Laos, Thailand, Malaysia,
2006). During these workshops, species conservation status Brunei, Vietnam, Cambodia and Indonesia (Kalimantan).
was reviewed in accordance to the IUCN red-list criteria ver- The datasets covered a total of 351 species and were kindly
sion 3.1 (Table 1; IUCN, 2001). provided by C. Francis (unpublished data), R. Steinmetz
(unpublished data), R. Boonratana (unpublished data), G.
2.1. Distribution models Csorba (unpublished data), J. MacKinnon (unpublished data),
D. Lunde (Lunde et al., 2003), E. Meijaard (unpublished data),
We considered 1077 species (99.2% of the total 1086 species in and J. Walston (unpublished data).
the database) in all the subsequent analyses. We excluded We integrated these existing datasets with point data on
four introduced species and five species with no reliable infor- species presence directly collected in the field in Indonesia,
mation on their distribution. For 901 species, we built deduc- The Philippines and Vietnam. We selected these three coun-
tive distribution models (DM; sensu Corsi et al., 2000) using the tries for several reasons including available logistic support,
available species–habitat relationships and environmental political stability, security for the field crew, cost of expedition
layers. First of all, land-cover and elevation were reclassified and permission to conduct research activities. The field crews
following the available suitability scores (see paragraph above were composed by two researchers from the Institute of Ap-
for more details on the definition of the scores). Then, for plied Ecology (IEA) and by two zoologist from the Indonesian
each species we combined the suitability scores of the two Institute of Sciences for Indonesia, by one researcher from
layers into a synthetic suitability index (Table 2). IEA and by two zoologist from the Institute of Biology of the
For four strictly water dependent species (Aonyx cinerea, Lu- University of the Philippines–Diliman for the Philippines, by
tra lutra, Lutra sumatrana, Lutrogale perspicillata) the habitat two researchers from IEA and by two zoologists from the
suitability scores were computed inside a 3-km buffer around Institute of Ecology and Biological Resources of the Vietnam-
water bodies and courses. For 63 species no information on ese Academy of Science and Technology in Vietnam. In each
the elevation range was available; in this case the suitability country, we followed a systematic random sampling design
was calculated considering only land-cover. to select 100 points (for a total of 300 points). The field crews
We defined as area of potential species presence the area reached the village that was closest to each random point and
inside the EOO that was classified as highly suitable by the conducted a direct and standardized interview (Boitani et al.,
DM. We did not build a DM for 144 species with EOO < 1000 km2 1999) with local hunters and/or villagers (1–5 villagers/hunt-
(too small an extent compared with the extent of our study ers actively participating in the interviews) aimed at collect-
area) and for other 32 species that had unreliable species– ing information on the current presence of 148 species of
habitat relationships (i.e. species whose ecology was not medium and large sized mammals (we considered species
known). For these 176 species, we used their entire EOO as the easier to recognize and of greater economic value for the local
 B I O L O G I C A L C O N S E RVAT I O N
populations) in the immediate neighborhood of the point.
Interviews were conducted in the local language with support
of pictures of the species; the ability of the interviewees in
recognizing species was tested using dummy pictures of com-
parable European and/or African species. Only (subjectively
judged) reliable information was retained. Not all 300 points
were reached by the field crews because of logistic reasons:
70 out of the original 100 points were sampled in Indonesia
and in the Philippines, 80 out of the original 100 points were
sampled in Vietnam, for a total of 220 sampled points freely
accessible through SAMD’s website.
Merging all datasets, we obtained points of presence for
399 species. To decrease spatial dependence, we removed all
locations closer than 5 km one from each other from each
species dataset. Moreover, we excluded all species with less
than five presence points from the validation analyses,
obtaining a final list of 190 species (21% of all the DMs) for
which model validation was possible.
To account for possible location errors and the spatial
uncertainty naturally associated with each point of presence,
we built a circular buffer (1.5 km radius) around each point of
presence (thus a total of nine grid cells was considered for
each point). A point of presence and the corresponding DMs
were considered to agree if at least one of the nine cells was
classified as highly suitable. In this way, for each species we
calculated the percentage of presence points that agreed with
the DM.
To test the significance of the agreement between points of
presence and the DMs, we used a permutation test following
Maiorano et al. (2007). We compared the percentage of agree-
ment calculated for the points of presence with that obtained
with 1000 sets of random points sharing the same character-
istics as the set of points of presence (same number of points,
distance among points equal or greater than 5 km). If the per-
centage of agreement calculated for the points of presence
was in the top 5% of the agreements obtained from the 1000
random samples, the model was considered validated.
To test whether the 190 species for which the points of
presence were available were representative of the ecology
of all 901 species with a DM, we compared the distribution
of the suitability scores among the two groups. In particular,
for each land-cover class, we calculated the percentage of
the 190 species for which the class represented a suitable
habitat; we calculated the same percentage for the 711 spe-
cies with a DM but without validation points, and we com-
pared the two distributions using a Kolmogorov–Smirnov test.
2.3. Potential species richness
We used the 901 DMs and the 176 EOOs to build a map of po-
tential species richness, calculated as the sum of all the spe-
cies’ areas of potential presence (i.e. for species with a DM the
areas inside the EOO classified as suitable, or the entire EOO
for species without a DM as defined in the Distribution models
section above). Moreover, to highlight the areas with a high
concentration of endemic or restricted-range species, we also
built a map of potential species richness in which each spe-
cies was weighted according to the inverse of its area of po-
tential presence (hereafter called potential weighted
richness). To highlight the areas with a high concentration
1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2733
of threatened species, we built a map of potential species
richness for threatened species considering only those spe-
cies classified as critically endangered, endangered, vulnera-
ble, or near threatened following the IUCN red-list criteria
version 3.1. The same maps of potential species richness were
calculated considering the EOOs only for all 1077 species.
2.4. Gap analysis
We performed two gap analyses: one considering a combina-
tion of the 901 DMs and 176 EOOs, and one considering EOOs
only. Gap analysis requires the identification of a representa-
tion target (Scott et al., 1993). We used a species-specific repre-
sentation target depending on the area of potential presence
for each species. We performed our analyses following the
representation target defined in the ‘‘global gap’’ project
(Rodrigues et al., 2004a): species with a narrow distribution
(area of potential presence smaller than 1000 km2) should be
protected in 100% of the area of potential presence; wide-
spread species (area of potential presence greater than
250,000 km2) should be protected in 10% of the area of poten-
tial presence; species with area of potential presence greater
than 1000 km2 and smaller than 250,000 km2 have a represen-
tation target that is obtained interpolating between the two
extremes using a linear regression on the log-transformed
area of potential presence.
A species not represented at all in any PA was considered a
total gap, a species whose representation target is only par-
tially met was considered a partial gap, and a species whose
representation target is met was considered covered.
3. Results
PAs do not cover the countries in the study area with similar
proportions (Table 3). For example, more than 44% of Brunei
Darussalam and less than 6% of Singapore is protected. PAs
cover a disproportionate percentage of mountainous area,
the median elevation inside PAs being 438 m (interquartile
range = 753 m) while the median elevation inside the study
area is 190 m (interquartile range = 516 m).
3.1. Validation
The 190 species that we considered in the validation proce-
dure were representative of the distribution patterns of all
species. We found no significant difference (p = 0.95) between
the distribution of the high suitability scores for the 190 spe-
cies and those for the 711 species that were not processed for
validation (Fig. 1). The main difference (although not statisti-
cally significant) was for the land-cover class ‘‘Tree Cover,
Regularly Flooded, Fresh Water’’, for which the 190 species
with validation show a higher percentage of high suitability
scores compared to the entire species dataset.
On average, 32 presence points (median = 20) were avail-
able for the 190 species with a maximum of 233 points for Par-
adoxurus hermaphroditus and a minimum of five points for
nine species (Babyrousa babyrussa, Sus cebifrons, Hipposideros
rotalis, Myotis annectans, Phoniscus atrox, Presbytis chrysomelas,
Pygathrix nigripes, Hylobates agilis, Hylopetes nigripes). For 140
out of 190 species (73.7%), the DMs gave positive validation re-
2734 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4

Table 3 – Country area, percentage being protected, and number of protected areas in each Southeast Asian country
Country Country area (km2) % In protected areas Number of protected areas

Brunei Darussalam 5898 44.25 44

Cambodia 182,062 24.02 30
Indonesia 1,911,253 14.10 431
Lao PDR 230,662 16.22 25
Malaysia 331,165 16.88 516
Myanmar 669,660 6.31 48
Papua New Guinea 397,160 11.43 35
Philippines 296,940 10.81 198
Singapore 592 5.57 6
Thailand 516,906 20.04 203
Viet Nam 328,809 5.96 99

Fig. 1 – Distribution of the high suitability scores for the species with a distribution model. Land-cover class: 1 – Tree Cover,
Broadleaved, Evergreen; 2 – Tree Cover, Broadleaved, Deciduous, Closed; 3 – Tree Cover, Needle-Leaved, Evergreen; 4 – Tree
Cover, Regularly Flooded, Fresh Water; 5 – Tree Cover, Regularly Flooded, Saline Water; 6 – Mosaic: Tree Cover/Other Natural
Vegetation; 7 – Shrub Cover, Closed-Open, Evergreen; 8 – Shrub Cover, Closed-Open, Deciduous; 9 – Herbaceous Cover,
Closed-Open; 10 – Sparse Herbaceous Or Sparse Shrub Cover; 11 – Cultivated and Managed Areas; 12 – Mosaic: Cropland/Tree
Cover/Other Natural Vegetation; 13 – Mosaic: Cropland/Shrub and/Or Grass Cover; 14 – Bare Areas; 15 – Water Bodies; 16 –
Artificial Surfaces.

sults, which were statistically significant for 46 species
(24.2%) at the a = 0.05 level, and for 77 species (40.6%) at the
a = 0.1 level. For 50 species (26.3%), the percentage of agree-
ment among DMs and points of presence was lower than
the agreement obtained with 1000 sets of random points,
but only for eight species (4.2%) the difference was signifi-
cantly lower (p < 0.05). We found no taxonomic bias in the
validation results, with the exception of Chiroptera and Pri-
mates. Considering the proportion of Chiroptera species over
the total sample of species with validation points, we found a
percentage of positively validated species greater than ex-
pected. On the contrary the validation results for Primates
showed a percentage of species with a negative validation
that was greater than expected.
3.2. Potential species richness
In general, the richness maps estimated using only EOOs were
similar to those estimated using both EOOs and DMs, although
the former maps depicted a much coarser spatial pattern. The
paragraphs below focus on describing potential species rich-
ness maps estimated using both EOOs and DMs, and point
out noteworthy differences with the maps calculated using
only EOOs. All potential species richness maps calculated
using the EOOs only are available in Appendix 1 online.
Considering potential species richness, the richest areas
(Fig. 2a) are found in Borneo (mainly Sarawak and Sabah),
Western Sumatra, Annamites mountains, and Malay Penin-
sula (Malaysia). Other important areas of high potential spe-
cies richness were also found in the Cardamom mountains
in Cambodia and Myanmar, along the mountain ranges of Pa-
pua New Guinea, and in Sulawesi. The relative importance of
Papua New Guinea and of Sulawesi was lower if considering
potential species richness calculated with EOOs only (Fig. 1a
in Appendix 1).
The map of potential weighted richness (Figs. 1b and 2b in
Appendix 1) showed a different pattern, with higher values
being concentrated mainly along mountain ranges (rough-
ly > 500 m a.s.l.) and in small islands. In particular, the richest
areas were found in the Annamite mountains (Vietnam and
Laos) and in the far north of Myanmar, which was the most
important area for weighted species richness in Indochina.
 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2735

Fig. 2 – (a) Potential species richness calculated as the sum of each species’ area of potential presence (EOO or DM); (b)
potential weighted species richness calculated as the sum of each species’ area of potential presence (EOO or DM): each
species is weighted according to the inverse of its area of potential presence; (c) potential threatened species richness
calculated as the sum of each species’ area of potential presence (EOO or DM) considering only critically endangered,
endangered, vulnerable or near threatened species. All maps are represented using a histogram equalize stretch in ArcGis;
the legends of (a) and (c) report the actual ranges of potential richness values, the legend of (b) reports percentages.
2736 B I O L O G I C A L C O N S E RVAT I O N
Fig. 2 (continued)
The Malay peninsula (Malaysia) was also a very important
area, as well as the Philippines (especially Palawan, Mindoro,
and Luzon), Sulawesi, New Guinea and a number of small is-
lands (Moluccas, Singapore, Natuna Besar, Mentawai, Engg-
ano, Simeleu, Sangihe, Talaud, Peleng, Waigeo, and Geelvink
Bay islands).
The map of potential richness for threatened species (Figs.
1c and 2c in Appendix 1) clearly shows the importance for con-
servation of areas like the Annamite mountains, Borneo (Sara-
wak in particular), Western Sumatra and peninsular Malaysia.
3.3. Gap analysis
Considering a combination of DMs (901 species) and EOOs
(176 species), only 88 species (8.2%) of the Southeast Asian
mammals that we considered were total gap. This number re-
mained almost unchanged considering EOOs only, with 81
species (7.5%) being considered total gap (as it was expected
given that 72% of the 88 total gap species above do not have
a distribution model). Considering both EOOs and DMs, the
orders with the highest numbers of total gap species were
the Rodentia (37 species) and the Chiroptera (28 species). Per-
amelemorphia, Dasyuromorphia, and Artiodactyla showed
relatively low numbers of total gap species (respectively
two, two, and six species), but extremely high percentages,
with Peramelemorphia being the order with the highest per-
centage of total gap species (Table 4). Considering EOOs only,
the results were almost unchanged, with the exception of
Dasyuromorphia (for which no species was classified as total
gap according to the EOO only analyses) and Artiodactyla
(that passed from five total gap species considering EOOs
and DMs to three species with EOOs only).
1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4
Considering a combination of EOOs and DMs, more than
59% of all the species (636 species) were partial gap species
and 33% (352) were fully covered (Table 4). The taxa with the
highest percentages of partial gap species were Soricomor-
pha, Rodentia, Diprotodontia, Lagomorpha and Primates,
but many other taxa showed high numbers of partial gap spe-
cies (Table 4). Considering EOOs only, 51.6% of all the species
(556 species) were partial gap species and 41% (440) were fully
covered (Table 4). Once more, Soricomorpha, Rodentia, Diprot-
odontia, Lagomorpha and Primates, although with different
percentages, were the taxa with the highest percentage of
partial gap species. The complete list of total and partial
gap species is available in Appendix 2 available online.
Considering threatened species (i.e. those species classi-
fied as critically endangered, endangered, vulnerable, or near
threatened according to the IUCN criteria), from 73.9% (EOOs
plus DMs) to 68.2% (EOOs only) are classified as total or partial
gap (Table 4), with Dasyuromorphia (only when considering
EOOs plus DMs), Diprotodontia and Rodentia being the taxa
with the highest percentages of threatened species not cov-
ered by any PA, and Erinaceomorpha, Peramelemorphia, and
Soricomorpha being the taxa with the highest percentages
of threatened species only partially covered by PAs. Notable
are also the cases of the Dermoptera, Lagomorpha and Probo-
scidea that, considering threatened species, are completely
covered by existing PAs (Table 4).
The areas with the highest numbers of total and partial
gap species (Fig. 3a) are Sulawesi, Papua New Guinea, and
Mentawai islands, together with many mountainous areas
(Annamite mountains at the boundaries between Laos and
Vietnam, peninsular Malaysia, North Myanmar, Luzon in
the Philippines, and northern Borneo). The same distribution
 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2737

Table 4 – Percentage of total gap species, partial gap species, and covered species for each order calculated using a
combination of DMs and EOOs

Order Total gap species (%) Partial gap species (%) Covered species (%)
All species Threatened species All species Threatened species All species Threatened species

Artiodactyla 9.6 (5.8) 3.8 (0.0) 51.9 (51.9) 63.8 (61.8) 38.5 (42.3) 32.4 (38.2)
Carnivora 1.9 (1.9) 0.0 (0.0) 29.6 (18.5) 20.8 (16.7) 68.5 (79.6) 79.2 (83.3)
Chiroptera 8.5 (7.9) 4.1 (4.1) 57.9 (48.8) 79.5 (72.6) 33.5 (43.3) 16.4 (23.3)
Dasyuromorphia 12.5 (0.0) 33.3 (0.0) 50.0 (50.0) 33.3 (66.7) 37.5 (50.0) 33.3 (33.3)
Dermoptera 0.0 (0.0) 0.0 (0.0) 50.0 (50.0) 0.0 (0.0) 50.0 (50.0) 100.0 (100.0)
Diprotodontia 6.9 (6.9) 13.3 (13.3) 67.2 (55.2) 76.7 (70.0) 25.9 (37.9) 10.0 (16.7)
Erinaceomorpha 0.0 (0.0) 0.0 (0.0) 57.1 (57.1) 100.0 (100.0) 42.9 (42.9) 0.0 (0.0)
Lagomorpha 0.0 (0.0) 0.0 (0.0) 66.7 (66.7) 0.0 (0.0) 33.3 (33.3) 100.0 (100.0)
Monotremata 0.0 (0.0) 0.0 (0.0) 50.0 (50.0) 33.3 (33.3) 50.0 (50.0) 66.7 (66.7)
Peramelemorphia 16.7 (16.7) 0.0 (0.0) 50.0 (41.7) 100.0 (100.0) 33.3 (41.7) 0.0 (0.0)
Perissodactyla 0.0 (0.0) 0.0 (0.0) 33.3 (33.3) 33.3 (33.3) 66.7 (66.7) 66.7 (66.7)
Pholidota 0.0 (0.0) 0.0 (0.0) 33.3 (33.3) 33.3 (33.3) 66.7 (66.7) 66.7 (66.7)
Primates 6.6 (6.6) 8.1 (8.1) 63.2 (52.6) 66.1 (54.8) 30.3 (40.8) 25.8 (37.1)
Proboscidea 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 100.0 (100.0) 100.0 (100.0)
Rodentia 9.7 (9.4) 13.2 (12.3) 63.1 (57.9) 69.3 (68.4) 27.2 (32.7) 17.5 (19.3)
Scandentia 5.9 (5.9) 9.1 (9.1) 35.3 (23.5) 36.4 (18.2) 58.8 (70.6) 54.5 (72.7)
Soricomorpha 5.4 (5.4) 0.0 (0.0) 76.8 (64.3) 100.0 (100.0) 17.9 (30.4) 0.0 (0.0)
All orders 8.2 (7.5) 8.1 (7.3) 59.1 (51.6) 65.8 (60.9) 32.7 (40.9) 26.1 (31.8)

Results obtained considering EOOs only are given in parenthesis. Threatened species include critically endangered, endangered, vulnerable,
and near threatened. The percentages are calculated considering the number of species in Table 1.

pattern was found considering total and partial gap species
potential richness calculated using EOOs only (Fig. 2a in the
Appendix 1), with the exceptions of North Vietnam and Min-
danao (Philippines), whose potential species richness was
higher.
Total gap species were distributed throughout the study
area (Fig 3b), from the extreme north of Myanmar to the SE
coast of Papua New Guinea. Many small islands (among
which Tawitawi, Dinagat, and Camiguin in the Philippines,
Natuna Besar, Simeleu, Sangir, and Peleng in Indonesia, Obi,
Kai, and Aru among the Moluccas) hosted total gap species.
A particularly important case is in the Mentawai archipelagos
(particularly Sipura and North Pagai), west of Sumatra, where
the highest number of total gap species (five species) was
found. Considering total gap species richness calculated
using EOOs only (Fig. 2b in Appendix 1), the general pattern
is almost the same, with the main exceptions of Central Prov-
ince in Papua New Guinea and of the mountain range be-
tween Irian Jaya and Papua New Guinea where no total gap
species was present.
The areas with the highest number of partial and total gap
species classified as threatened (Fig. 3c) are mainly located in
Sulawesi, in Malaysian Borneo, and in the Mentawai islands.
Considering mainland Southeast Asia, the Annamite Moun-
tains and the Malay Peninsula (Malaysia) have a particularly
important role. Also in this case, the potential richness pat-
tern obtained considering EOOs only was almost the same
(Fig. 2c in Appendix 1), with higher richness values in West
Java and in Mindanao, Negros and Mindoro (Philippines).
Most of the species whose conservation target was com-
pletely met have a relatively large distribution range (Fig. 4a).
On the contrary, total gap species (Fig. 4b) have a small to
medium distribution range, a distribution which is similar to
that of the species completely enclosed in PAs (Fig. 4d).
4. Discussion
Our study represents an important contribution to mammal
conservation in Southeast Asia. It provides a new and com-
plete dataset on all mammal species of the region and pre-
sents a synthetic view of the conservation status in relation
to the network of existing PAs. Several multi-species gap anal-
yses have been carried out globally (e.g. Rodrigues et al.,
2004a) or on a sub-continental scale (e.g. Fjeldså et al., 2004;
De Klerk et al., 2004) but no comprehensive gap analysis has
ever been performed for the Southeast Asian region. More-
over, extensive modeling of species distribution has been ap-
plied at national level (e.g. Maiorano et al., 2006) or for other
continents (e.g. Rondinini et al., 2005), but no systematic ef-
fort has ever been applied to the Southeast Asian region.
Many international conservation efforts have considered our
study area (McNeely et al., 1990; Olson and Dinerstein, 1998;
Myers et al., 2000; Wikramanayake et al., 2002) and many dif-
ferent PAs have been established (currently more than 1700
according to the WPDA, 2006). Although the effectiveness of
networks of PAs in protecting biodiversity is often debated
and different studies have produced different results (e.g.
Bruner et al., 2001 vs. Sodhi et al., 2004a), in Southeast Asia
the contribution of PAs to conservation of an important taxo-
nomic group such as mammals has never been tested. Our
analysis clearly indicates that the existing PAs are inadequate
in assuring the conservation of mammals across the region.
We based our results both on deductive distribution mod-
els and on extents of occurrence, with a few discrepancies
among the two analyses as already discussed in Rondinini
et al. (2005) and Loiselle et al. (2003). Deductive distribution
models have been successfully used elsewhere (Rondinini
et al., 2005; Maiorano et al., 2006, 2007) reducing the level of
commission errors that is naturally present in species distri-
2738 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4

Fig. 3 – (a) Potential gap species richness calculated as the sum of each species’ area of potential presence (EOO or DM)
considering only total and partial gap species; (b) potential presence of total gap species (EOO or DM); (c) potential threatened
gap species richness calculated as the sum of each species’ area of potential presence (EOO or DM) considering only total and
partial gap species that were classified as critically endangered, endangered, vulnerable or near threatened. Maps a and c are
represented using a histogram equalize stretch in ArcGIS and report the actual ranges of potential richness values. The list of
total and partial gap species can be found in the Appendix 2 available online.
 B I O L O G I C A L C O N S E RVAT I O N
Fig. 3 (continued)
bution maps (Guisan and Zimmermann, 2000; Scott et al.,
2002; Loiselle et al., 2003; see Jetz et al. 2008 for an analysis
of the ecological correlates). However, there is no inherent
assurance that a distribution model portrays reality (Guisan
and Zimmermann, 2000; Johnson and Gillingham, 2004) and
a model that poorly represents the presence of a species
may result in harmful conservation and management actions
(Loiselle et al., 2003; Wilson et al., 2005). Therefore, model val-
idation is an essential step in any application of models to
conservation, and it is an important component of our study.
We were able to perform a validation analysis for only 21% of
all the distribution models that we developed (190 out of 901
distribution models), but we were able to cover a wide taxo-
nomic range (12 out of 17 orders), a wide geographic range
(from Thailand to Philippines/Sulawesi), and we represented
the distribution patterns of all mammal species. Moreover,
the results of the validation process can be considered to be
positive. The distribution models predicted potential pres-
ence significantly better than random at the a = 0.05 for
22.2% of the species, and for only 4.2% of the species were
the models significantly worse than random at the same level
of significance. For all other distribution models, no statisti-
cally significant result was found, either for the large percent-
age of suitable areas in the models or the low number of
available points of presence.
However, raising the significance level to a = 0.1, the per-
centage of distribution models which portray species distri-
bution worse than random was almost stable (6.3%), while
almost 41% of the models can be considered to be signifi-
cantly better than random. We can conclude that our models
represent a reasonable baseline for conservation planning at
the scale of our study area.
1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2739
We found no particular bias in the validation results with
reference to the different taxonomic groups. The only excep-
tion were the Chiroptera, for which we obtained better valida-
tions than expected, and the Primates for which the
proportion of validated species was lower than expected.
Clearly, the positive results obtained for Chiroptera are linked
to the higher quality of the validation points; for almost all
the species of Chiroptera considered for validation, we ob-
tained presence locations from data collected in the field
(mainly captures and scientific observations). Considering
Primates, most of the validation points were of lower quality
(if compared with those available for Chiroptera), being col-
lected mainly through interviews and often located close to
small patches of suitable habitat (observed directly in the
field) that are almost invisible at the scale of our distribution
model.
We have provided no measure of commission or omission
error associated to our distribution models. Both types of er-
rors are important in any conservation planning exercise,
with omission errors that affect the comprehensiveness of a
network of PAs, and commission errors that affect represen-
tativeness and adequacy of a reserve network (Rondinini
et al. 2006a). No direct measure of the commission error rate
was possible with out dataset, and yet it has been argued that
conservation decision makers should prefer models that min-
imize commission errors because such errors lead to the
selection of reserves that do not actually contain the target
species (Loiselle et al. 2003; Rondinini et al. 2005; Jetz et al.
(2008)) have demonstrated that EOOs can overestimate the
real range occupancy (conditional on EOO size) with propor-
tions going from 0% to 91%, with an average of 39%. However,
Jetz et al. (2008) performed their analyses on birds in North
2740 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4

(a) area of potential presence x protection error rate cannot be considered negligible and that it may
100

10 have relevant consequences in gap analyses (Rondinini et

partial gap
covered al., 2006). However, we used two general approaches to esti-
80

mate the frequency of gap, partial gap, and covered species,

% protected

one using EOOs only and the other using a combination of

60

DMs and EOOs. The results of these two approaches should

40

5 provide a range of reasonable estimates that likely brackets

20 12 the true value of the parameter of interest. In fact, while
0
20

90
60
estimates based on EOOs only should minimize omission er-
10
5 rors at the cost of potentially high commission errors, esti-
30 30 20 10 5
0

1 2 3 4 5 6 7 mates based on a combination of DMs and EOOs should

log area of potential presence (km2) reduce commission errors at the expenses of higher omis-
sion errors.
number of species

(b) 0% protected
A further point regarding our distribution models and ex-
tents of occurrence calls for caution in the interpretation of
0 15

our results. We have produced (validated) estimates of habitat

1 2 3 4 5 6 7 suitability and reliable maps of distribution ranges but we do
log area of potential presence (km2) not have any insurance that the species are effectively pres-
ent in their entire EOO or even in the suitable part of their
number of species

(c) 20% protected

EOO only. This is the well-known problem of the ‘‘empty for-
0 75 150

est syndrome’’, with large animals (mainly primates, carni-

vores and ungulates) that are extinct in vast areas of their
former EOO because of commercial hunting, even if the vege-
1 2 3 4 5 6 7
log area of potential presence (km2) tation is still intact (Redford, 1992; Milner-Gulland et al., 2003;
Corlett, 2007). This can have profound influences on our re-
number of species

(d) 100% protected

sults, as it can be demonstrated considering the case of the
0 10 20

banteng (Bos javanicus) and of the Hose’s lead monkey (Presby-

1 2 3 4 5 6
log area of potential presence (km2)
Fig. 4 – (a) Number of species for each class of range size and
for each class of proportion of area of potential presence
inside protected areas (EOOs plus DMs). The grey area
represents the protection target we set for species. Isolines
enclose areas with equal numbers of species (dotted lines
620 species, continuous lines >20 species). (b) Number of
total gap species for each range size class. (c) Number of
species for each range size class with 20% of their area of
potential presence protected. This graph corresponds to the
peak number of species in (a). (d) Number of species for each
range size class with 100% of their area of potential
presence protected. Same results (not shown) were obtained
for EOOs only.
America, South Africa and Australia. We performed our anal-
yses on mammals in South East Asia, with a much smaller le-
vel of knowledge on species ecology and distribution,
particularly for Rodentia, Soricomorpha, and Chiroptera that
make up 71% of our sample. Thus we can assume that the
commission error associated with the distribution models is
much smaller if compared to the commission error associated
to the available EOOs, probably with a proportion that is
greater than that measured by Jetz et al. (2008).
The omission error rate can be easily obtained from the
percentages of agreement that we calculated during the val-
idation procedure. In particular, an average 32% (standard
deviation = 21%) of the occurrence points were not covered
by suitable areas in the DMs. We recognize that the omission
tis hosei). According to our results (see the Appendix 2 and
consider the distribution model) the representation target
7
for the banteng is more than met (123% of the representation
target is covered by PAs). However, the species is gone from
most of the highly suitable areas (even though Java still repre-
sent one of the species’ stronghold), mainly because of illegal
hunting for the trade in horns (Hedges, 2000; Steinmetz, 2004).
In the case of the Hoses’s leaf monkey the representation tar-
get is almost completely met (87% according to our results)
but a recent report (Nijman, 2005) outlines that hunting for
medicinal bezoar stones have produced local extinction of
the species even in highly suitable areas as the huge and re-
mote Kayan Mentarang National Park in Indonesia. However,
our results are still extremely important, because we provide
at least an estimate of how much suitable habitat remains in
relation to the existing PAs, an estimate that suffers with low-
er commission errors if compared to the original EOO (the
representation target met as measured using the EOOs is
180% for the banteng and 130% for the Hoses’s leaf monkey)
and that is important for planning from the standpoint of po-
tential recovery for populations.
As in any gap analysis, our results heavily depend on the
dataset and on the pre-determined representation targets.
In particular, even though we applied a representation target
that has been used in the previous studies, we are not explic-
itly accounting for species viability (Svancara et al., 2005).
Even considering all the inherent limitations and uncer-
tainties that characterize analyses on a large scale and our
dataset, our results give a clear indication of important pat-
terns in the current distribution of mammals in Southeast
Asia. The analysis of potential species richness demonstrates
that many large areas are still potentially occupied by a high
number of species, even though many of the flat areas, where
 B I O L O G I C A L C O N S E RVAT I O N
most of the human pressure is concentrated, are relatively
poor in mammal species. Particular caution is necessary in
considering the results on potential weighted species rich-
ness. In fact, the areas of high species richness located at
the boundaries of our study area are probably influenced by
the presence of species that have a much wider distribution
outside of our study area but that seem to have a small range
when considering only the zone within our study area. For
example, this is the case of the tufted deer (Elaphodus cephalo-
phus), a total gap species, and of the red panda (Ailurus ful-
gens): both species occupy a tiny range in northern
Myanmar, but outside of our study area the first is distributed
throughout central and southern China and the latter occurs
in Nepal, India, Bhutan, and South China. However, the areas
with high potential weighted richness in the smaller islands
and in the central parts of our study area are clearly an indi-
cation of important centers of restricted species ranges. For
example, at least 15 endemic species are present in the Men-
tawai archipelago, with one Chiroptera species (classified as
data deficient) that is total gap, five threatened and gap Pri-
mates species (among which the total gap and critically
endangered Macaca pagensis and the partial gap and critically
endangered Simia concolor), eight threatened and gap Rodentia
species (among which the total gap and endangered Hylopetes
sipora), and the endangered and total gap Tupaia chrysogaster.
We have also shown that an important number of species
are not covered by any PAs. These species are distributed
throughout the entire study area and their conservation calls
for the creation of new PAs that can contribute to filling the
existing conservation gap. This would be particularly impor-
tant for total gap species over small islands, where species
survival is often endangered by the presence of introduced
species as well as habitat vulnerability and other factors (Pur-
vis et al., 2000; Sodhi et al., 2004b).
Considering gap species, many small semi-natural areas
surrounded by cultivated areas, both in the mainland and in
the larger islands, should be considered for the establishment
of new PAs or, better, for the implementation of effective
management plans, but clearly Papua New Guinea (where
many total gap species occur), Sulawesi, and the internal
mountainous areas represent the areas with the highest con-
servation priority (Fig. 3). Particular attention should be
placed towards species like the saola (Pseudoryx nghetinhensis),
a highly distinctive monotypic genus first described 15 years
ago (Dung et al., 1993) that is highly endangered because of
habitat loss and hunting (Timmins et al., 2007): almost half
of its range is covered by PAs but all the necessary efforts
should be made to extend protection to the entire range of
the species along the Annamite mountains (Vietnam–Laos
borders) where the last know populations of the species occur
at extremely low densities.
Our results are comparable in some ways to those ob-
tained by Rodrigues et al. (2004a), who identified some of
the most important areas for the conservation of mammals
in Southeast Asia. Rodrigues et al. (2004a) developed two
possible global scenarios and found that 5.5–11% of all mam-
mal species were not covered by any PA, a percentage that is
similar to our 8.2–7.5% of total gap species. However, for par-
tial gap species, our results are different: using the same
representation target, Rodrigues et al. (2004a) found that
1 4 1 ( 2 0 0 8 ) 2 7 3 0 –2 7 4 4 2741
34% of the species were partial gap, while we found that
52–59% of the species were partial gap. Our results were ob-
tained using both distribution models and EOOs, while
Rodrigues et al. (2004a), besides considering a different study
area and working on a different scale, used species’ EOOs
only, introducing a greater proportion of commission errors
in their analyses (Rondinini et al., 2005). This difference
clearly explains the smaller percentage of partial gap species
(Loiselle et al., 2003; Rondinini et al., 2005) and provides an
indication that our results might be more reliable for conser-
vation applications.
The results we obtained for partial gap species are obvi-
ously important for the identification of areas in need of fur-
ther attention from the conservation community, but the
results we obtained for total gap species are particularly
important. In fact, we found a relatively low number of spe-
cies that are total gaps (8.2–7.5% of the species). Yet a con-
siderably higher number of mammal species of the region
are classified as threatened according to the IUCN red-list
criteria. In fact, 34% of all 1086 species considered in the
SAMD database are listed in the IUCN red-list categories as
critically endangered, endangered, vulnerable, or near threa-
tened, while 45% are classified as least concern and 21% as
data deficient or not evaluated, thus being potential candi-
dates for a threatened category (red-list category assess-
ments reviewed during the five workshops; note that not
all the species assessments have already been officially con-
firmed by IUCN).
This is clearly an indication that PAs cannot be considered
as the ending point of our conservation strategies. Defining
PAs is an option that can be pursued relatively easily in many
situations. However, managing effective PAs for conservation
is much more difficult and expensive than just establishing
them and different studies have indicated that PAs in the tro-
pics usually act as ‘‘paper parks’’ (Schwartzman et al., 2000;
Curran et al., 2004; Fuller et al., 2004; Sigel et al., 2006; Sodhi
et al., 2006b; Verburg et al., 2006; Gaveau et al., 2007; but see
Bruner et al., 2001; Nagendra et al., 2004; Nepstad et al., 2006).
Moreover, for a number of species PAs do not ensure prop-
er conservation (Corlett, 2007). Notable examples can be
found among many different taxa, going from the Asian ele-
phant (Elephas maximus), to rhinos (Dicerorhinus sumatrensis
and Rhinoceros sondaicus), orangutans (Pongo pygmaeus and
Pongo abelii), pangolins (Manis spp.) to many other taxa (Cor-
lett, 2007) for which PAs have not been able to stop population
declines.
In this context, it is clear that the role of existing PAs, as
well as the establishment of new PAs, should be considered
in a broader framework of all local ecological and socio-eco-
nomic trends, including the growing human population,
growing economy and infrastructure development. PAs alone
cannot be the solution to all conservation problems. In fact,
we should also focus on off-reserve management and the
preservation of natural processes.
Acknowledgements
A great number of people and organizations supported the
project throughout its implementation. It is impossible to list
2742 B I O L O G I C A L C O N S E RVAT I O N
them all, but we would like to mention at least the following
individuals and organizations that provided data, collabora-
tion, criticism, review and general support: G. Amori, P. T.
Ahn, K. Aplin, D. Balete, P. Bates, R. Boonratana, L. X. Canh,
D. Cesarini, W. Duckworth, C. Francis, C. Groves, T. Kingston,
L. Heaney, K. Helgen, M. Hobbelink, M. Hoffman, D. Lunde, J.
MacKinnon, I. Marzetti, A. Montemaggiori, E. Meijaard, M.
Pedregosa, H. Q. Quynh, G. Reggiani, S. Roberton, M. Rulli, L.
Ruedas, V. Salvatori, S. Savini, W. Sechrest, M. Sinaga, R. Tim-
mins, N. Van Strien, J. Walston, Sapienza Università di Roma,
the Asean Regional Centre for Conservation of Biodiversity,
the University of the Philippines–Diliman, the Indonesian
Institute of Sciences, the Vietnamese Institute of Ecology
and Biological Resources. Richard Corlett and three anony-
mous reviewers provided helpful comments that greatly im-
proved our original manuscript.
Appendix A. Supplementary data
Supplementary data associated with this article can be found,
in the online version, at doi:10.1016/j.biocon.2008.08.019.
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