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Abstract

We highlight hitherto unreported populations of two globally threatened Phalangeridae species on southeast Sulawesi’s offshore islands – Bear Cuscus Ailurops ursinus and Small Sulawesi Cuscus Strigocuscus celebensis – and observations of a third range-restricted species – Peleng Cuscus Strigocuscus pelengensis. Our data are based on records made during 11 years of seasonal surveys on Buton, and short-term expeditions to Kabaena and Manui. Our observations of S. celebensis on Buton, where it occurs in three protected areas, represent an important range extension for this species, as do our observations of A. ursinus on Kabaena, where it is also widespread. We also report the unexpected presence of S. pelengensis on Manui. Buton in particular appears to be an important stronghold for both A. ursinus and S. celebensis, given that forest ecosystems here remain extensive and relatively intact. Both these species may also display a previously unreported adaptability to disturbed forest and even some non-forest habitats within our study area. Hunting pressures, a proven threat to these species in northern Sulawesi, may also be lesser here.
Distribution and status of threatened and endemic marsupials
on the offshore islands of south-east Sulawesi, Indonesia
Thomas E. Martin
A,I
, Joseph Monkhouse
A
, Darren P. OConnell
B
, Kangkuso Analuddin
C
,
Adi Karya
C
, Nancy E. C. Priston
D
, Charlotte A. Palmer
A
, Barnabas Harrison
E
, Jack Baddams
A
,
Abdul H. Mustari
F
, Philip M. Wheeler
G
and David G. Tosh
H
A
Operation Wallacea, Wallace House, Old Bolingbroke, Lincolnshire, PE23 4EX, UK.
B
School of Natural Sciences, Trinity College, Dublin 2, Ireland.
C
Department of Biology, Faculty of Mathematics and Natural Sciences, Halu Oleo University of Kendari,
Southeast Sulawesi, Indonesia.
D
Centre for Ecology, Environment and Conservation, Oxford Brookes University, Gipsy Lane, Oxford,
OX3 0BP, UK.
E
5 Shackleton Court, 2 Maritime Quay, Isle of Dogs, London, E14 3QF, UK.
F
Department of Forest Resources, Conservation, and Ecotourism, Faculty of Forestry, Bogor Agricultural University,
Bogor, Jawa Barat, Indonesia.
G
School of Environment, Earth, and Ecosystem Sciences, Faculty of Science, Technology,
Engineering and Mathematics, The Open University, Walton Hall, Milton Keynes, MK7 6AA, UK.
H
Centre for Environmental Data Recording, National Museums of Northern Ireland, 153 Bangor Road,
Holywood, County Down, BT18 OEU, UK.
I
Corresponding author. Email: tom_martin_2010@yahoo.co.uk
Abstract. We highlight hitherto unreported populations of two globally threatened phalangerid species on south-east
Sulawesis offshore islands bear cuscus (Ailurops ursinus) and small Sulawesi cuscus (Strigocuscus celebensis)and
observations of a third range-restricted species Peleng cuscus (Strigocuscus pelengensis). Our data are based on records
made during 11 years of seasonal surveys on Buton, and short-term expeditions to Kabaena and Manui. Our observations
of S. celebensis on Buton, where it occurs in three protected areas, represent an important range extension for this species,
as do our observations of A. ursinus on Kabaena, where it is also widespread. We also report the unexpected presence
of S. pelengensis on Manui. Buton, in particular, appears to be an important stronghold for both A. ursinus and S. celebensis,
given that forest ecosystems here remain extensive and relatively intact. Both these species may also display a previously
unreported adaptability to disturbed forest and even some non-forest habitats within our study area. Hunting pressures,
a proven threat to these species in northern Sulawesi, may also be lesser here.
Additional keywords: cuscus, Indonesia, marsupial, Phalangeridae, population.
Received 13 November 2017, accepted 16 February 2018, published online 16 March 2018
Introduction
The biodiversity hotspot of Wallacea, Indonesia, is notable for
its high rate of endemism, particularly in mammals, and its mix
of fauna of both Asian and Australasian origin (Whitten et al.
2002; Myers 2003). Sulawesi, the largest landmass in Wallacea,
represents the westernmost extent of marsupial species in Asia
(Whitten et al.2002), where ve members of the Phalangeridae
occur (Helgen and Jackson 2015). Four species are endemic to
Sulawesi and its satellite islands (including the Sula Islands).
The remaining species, the ornate cuscus (Phalanger ornatus), is
found in both far-northern Sulawesi and the Moluccas (Helgen
and Jackson 2015). These endemic Sulawesi cuscuses represent
an important global conservation priority, with one species
Talaud bear cuscus (Ailurops melanotis)being considered
Critically Endangered and a top 25 EDGEspecies (Flannery and
Helgen 2016; Zoological Society of London 2017), two species
bear cuscus (Ailurops ursinus) and small Sulawesi cuscus
(Strigocuscus celebensis)being considered Vulnerable
(Helgen et al.2008; Salas et al.2008), and the nal species
Peleng cuscus (Strigocuscus pelengensis)being considered
Least Concern, but of very restricted geographic range (Leary
et al.2016a).
The Sulawesi Phalangeridae remain poorly studied (Helgen
and Jackson 2015). Little is known about their ecology, the
Journal compilation Australian Mammal Society 2018 www.publish.csiro.au/journals/am
CSIRO PUBLISHING
Australian Mammalogy
https://doi.org/10.1071/AM17052
threats they face, and even their distribution within Sulawesi
(although see Dwiyahreni et al.1999; Farida and Dahruddin
2008). Here, we report observations of three endemic cuscus
species on south-east Sulawesis offshore islands, made during
the course of 13 years of seasonal eldwork. We highlight
the existence of previously unreported populations of cuscus on
these islands, discuss the apparent status of these populations,
and provide a summary of ecological observations that will
improve existing knowledge of these poorly known marsupials.
Materials and methods
Study area
Fieldwork was focused on three islands off the coast of Sulawesis
south-eastern peninsula (Fig. 1): Buton, Kabaena and Manui. We
also briey visited the islands of Muna and Wowoni.
Buton, located off the mainlands south-eastern peninsula,
is the largest (~560 000 ha) of Sulawesis satellite islands. It is
mostly covered by rugged hills, with a few isolated peaks
reaching up to 1100 m. The island retains much of its original
natural vegetation cover, with 77% of its surface occupied by
forest (Martin et al.2015), and has been highlighted as an area
of high conservation value (Cannon et al.2007). Most of these
forested areas lie within formal protected areas, particularly
the 65 000-ha Lambusango Forest Reserve and the 810-ha
Kakenauwe Forest Reserve in the south of the island, and the
98 600-ha Buton Utara Forest Reserve in the north.
Kabaena, lying west of Buton, is the third-largest (~87 300 ha)
of south-east Sulawesis offshore islands. It is more mountainous
than its neighbouring islands, consisting of a single central
massif, Gunung Sambapolulu, which reaches 1560 m (Keim
2009). Forest cover here is patchy most lowland areas have
been cleared for agricultural land with pockets of rainforest
remaining in parts of the mountainous interior, along with more
extensive patches of savannah and open woodland (Gillespie
et al.2005). None of these remaining forest patches are formally
protected.
Manui is a small (~9000 ha), low-lying island (maximum
altitude 170 m) that is also the most biogeographically isolated
of south-east Sulawesis major satellites, located ~50 km east
of the mainland. It remains very poorly studied. It is covered
mostly in dry monsoon forest (a very different climax vegetation
compared with the seasonal rainforest found on the rest of
south-east Sulawesis offshore islands), farmland (primarily
cassava and coconut) and patches of rough scrub. No protected
areas exist on the island.
Muna, the second-largest of south-east Sulawesis offshore
islands (~289 000 ha) lies between Buton and Kabaena. It largely
comprises a low-lying (mostly <100 m) limestone plateau,
reaching a maximum elevation of 400 m (Milsom et al.1999).
This relatively at, easily accessible terrain has led to Muna
being almost entirely deforested and mostly covered with
plantations, arable farmland and scrubland, with only very small
pockets of forest remaining (Gillespie et al.2005). Wowoni
(~650 00 ha) lies north of Buton. Substantial tracts of forest
persist throughout much of the islands interior (Farida and
Dahruddin 2008); these are predicted to have high conservation
value (Cannon et al.2007), although its ecology remains largely
unexplored.
Survey work
These islands were visited as part of seasonal biodiversity
surveys run by Operation Wallacea (www.opwall.com). These
have run annually between June and August from 1996 to 2017,
with at least one author participating each year between 2004
and 2017 (except in 2015). Most eldwork has focused on
Buton, which was visited annually for eight weeks between
2004 and 2014. Kabaena was visited in JuneJuly 2016 for
four weeks, Manui was visited in August 2017 for one week,
and Muna and Wowoni were visited in July 2017 for one week
and two weeks respectively. Fieldwork on Buton, Kabaena and
Manui encompassed a range of forest and non-forest habitat,
while on Muna and Wowoni only farmland and scrub habitats
were visited.
No specic survey methodologies were employed to formally
assess cuscus populations on these islands; attempts to do so
failed, largely due to the difculty of detecting these species
using standardised methodologies. A. ursinus was frequently
observed in the forest and was technically a recordable species
in systematic mammal transect surveys; in practice, however,
these did not produce robust information on the species
occurrence, with encounters from these surveys averaging <1
observation per season. Fieldwork on Kabaena, Muna, Wowoni,
and Manui was focused specically towards an evolutionary
study of island birds. Therefore, cuscus observations on all
islands are the result of opportunistic encounters, pooled from
all authors, rather than from formal targeted surveys. To provide
supporting independently veried evidence for our observations,
we archived a research-grade (i.e. independently veried by at
least two other users) image of each cuscus species on each
island where they were observed on inaturalist (inaturalist.org).
Previously existing knowledge of speciesdistributions was
gathered from range maps produced by the IUCN (Helgen et al.
2008; Salas et al.2008; Leary et al.2016a), and by Helgen and
Jackson (2015), and information provided in Flannery (1995),
Suyanto et al.(2002) and Whitten et al.(2002).
Results
In total, we recorded three cuscus species on three of south-east
Sulawesis offshore islands. We discuss these observations in
the species accounts below. Links to our veried records can be
found in Table S1, available as supplementary material to this
paper. A selection of images is provided in Fig. S1.
Bear cuscus (Ailurops ursinus)
A. ursinus was observed on two islands, Buton and Kabaena
(Fig. S1a). It appears to be widespread and fairly common
on Buton, having been observed in three protected areas:
Lambusango Forest Reserve, Kakenauwe Nature Reserve, and
Buton Utara Nature Reserve. It has also been recorded in
multiple habitats, including primary and secondary forest, scrub,
and farmland. This includes one observation in July 2014 of an
individual found in an extensive area of degraded coastal scrub
(531018.900S, 12233055.200 E), with no connectivity to closed-
canopy forest ecosystems. It is difcult to detect within the
forest interior, normally being located high in the canopy.
However, it is often quite conspicuous in forest-edge ecosystems,
where high densities have been recorded. A 2.4-km roadside
BAustralian Mammalogy T. E. Martin et al.
survey detected a maximum of 15 individuals (6.25 individuals
per kilometre walked) in a single count; this was immediately
after an intense rainstorm.
The species was recorded at all three research locations
visited on Kabaena: Sikeli at 10 m (515048.200S, 12147045.200 E),
Enano at 132 m (515047.400 S, 12158008.600 E), and Tangkeno
at 650 m (516043.600S, 12155024.500E), suggesting it to be
widespread on the island. Observations at Enano and Tangkeno
were made in secondary forest, while at Sikeli a single
observation of an adult carrying a juvenile was made within an
(a)
(b)
Manui
Wowoni
Kabaena
Muna
Buton
40 20 0 40
Kilometres
N
Fig. 1. Map showing (a) the location of south-east Sulawesi (inset) within the Indonesian archipelago (total shaded
area), and (b) the locations of the offshore islands of Kabaena, Muna, Buton, Wowoni and Manui in relation to mainland
south-east Sulawesi.
Distribution of marsupials in south-east Sulawesi Australian Mammalogy C
area of highly degraded coastal farmland, with no connectivity
to surrounding forest patches.
In contrast to the ndings of Dwiyahreni et al.(1999), who
frequently observed A. ursinus in groups, most of our A. ursinus
sightings on both Buton and Kabaena were of single individuals
or a parent with a juvenile. Sightings of two adults together
were uncommon, and groups of more than three individuals
were observed very rarely.
Small Sulawesi cuscus (Strigocuscus celebensis)
S. celebensis was observed only on Buton (Fig. S1bd). It is
a cryptic species, given its nocturnal activity cycle (Whitten et al.
2002), and is difcult to survey systematically. Nevertheless,
it was encountered opportunistically relatively frequently,
typically being observed two or three times each eld season.
Most observations of this species were at night, with occasional
observations during the day of individuals disturbed by survey
work. These diurnal observations include two incidences of the
species being accidentally disturbed within, and subsequently
emerging from, hollow dead trees. The rst of these encounters
involved a single individual observed within a mixed coffee
and cashew plantation (511053.700S, 12250056.700 E) in July
2005, and the second involved an adult and juvenile observed
in closed canopy forest (511048.200S, 12253002.100 E) in July
2007. These records suggest that the species utilises these tree
hollows as sleeping sites during the day, as has been reported for
other nocturnal cuscus species (e.g. Flannery 1995; Salas 2002).
Observations were made throughout much of the islands
forested area, including within three protected areas:
Lambusango Forest Reserve, Kakenauwe Nature Reserve, and
Buton Utara Nature Reserve. Our observations of S. celebensis
have most frequently been made in closed canopy forest habitats,
although the species has occasionally been encountered in
edge forest, farmland, and within gardens near the city of Ereke
(448057.900S, 12310024.000 E) with no direct connectivity to
large tracts of closed canopy forest.
A further notable observation regarding S. celebensis records
from Buton relates to the variability in the colouration of
individuals. Helgen and Jackson (2015) describe the species as
being a uniform grey-brown colour dorsally, with a white
underbelly, while Flannery (1995) describes it as being drab
brown. On Buton we have observed individuals with dark grey,
grey-brown and sandy-brown dorsal fur (Fig. S1bd), and some
individuals have a black forehead stripe (Fig. S1b), which is
absent in others (Fig. S1c,d), suggesting that the speciespelage
is more variable than currently described.
Peleng cuscus (Strigocuscus pelengensis)
We recorded S. pelengensis on Manui (Fig. S1e,f), where two
separate individuals were observed on 7 August 2017 within
an extremely rocky, inaccessible area of dry forest growing over
aeld of large coral boulders (335043.000S, 12303009.900 E). The
rst individual was initially seen in a low bush growing between
these boulders; when disturbed, it crawled into a deep coral hole
and disappeared from view. The second individual was observed
in a tall tree. These individuals were readily distinguished from
S. celebensis by their noticeably larger and more heavily built
bodies, their orange-brown dorsal colouration and yellowish
belly fur, and their wider tail-base (Flannery 1995; Helgen
and Jackson 2015). They also lacked the distinctive dorsal stripe
of the larger-bodied P. ornatus (Flannery 1995; Helgen and
Jackson 2015).
Discussion
The results of our surveys on south-east Sulawesis offshore
islands identify important range extensions for all three cuscus
species detected specically, A. ursinus on Kabaena,
S. celebensis on Buton, and S. pelengensis on Manui. None of
our consulted sources indicate these species to be previously
recorded on these islands.
While our records of A. ursinus on Kabaena are new, it has
previously been recorded on Buton (Whitten et al.2002; Salas
et al.2008; Helgen and Jackson 2015). We did not record the
species on either Muna or Wowoni, although this might not
be surprising given our very short eldwork periods on these
islands. It is possible that the species persists on both these
islands, considering its presence on more biogeographically
isolated Buton and Kabaena, and as it has also successfully
colonised the Banggai and Togian Islands in eastern and
northern Sulawesi respectively (Flannery 1995; Salas et al.2008;
Helgen and Jackson 2015). On Muna it is considered to be
a formerly occurring species that may have been extirpated due
to habitat loss (Salas et al.2008). This may or may not be the
case. Small forest fragments remain on Muna that support
large-bodied mammals, such as the booted macaque (Macaca
ochreata) and the Sulawesi wild pig (Sus celebensis). These may
be sufcient to sustain a persistent population of A. ursinus,
given that the species appears to have some tolerance of non-
forest habitats on neighbouring islands (see below). It seems
likely that the species also occurs on Wowoni, given its relative
proximity to the mainland, the large tracts of forest ecosystems
remaining here, and because other cuscus species have been
shown to occur (Farida and Dahruddin 2008).
The fact that S. celebensis has not been previously reported
on Buton is somewhat surprising, given that the island has one
of the most extensive areas of suitable habitat for the species
anywhere in south-east Sulawesi. Our records of the species
presence in three reserves found on the island also partially
address a knowledge gap identied by Helgen et al.(2008)
regarding protected areas known to support this species.
Although we did not record S. celebensis elsewhere, Farida
and Dahruddin (2008) detected the species on Wowoni, an
important extension to the speciesknown range that has yet to
be incorporated into distribution maps.
Our records of S. pelengensis on Manui were unexpected,
given that the species is currently recorded only from the
Banggai and Sula islands, some 210 km to the north (Suyanto
et al.2002; Helgen and Jackson 2015; Leary et al.2016a). It is
unclear whether these individuals represent an established
population and whether they are from a natural colonisation
or anthropogenic introduction. In support of these being an
established, natural colonisation, the closely related S. celebensis
has colonised remote islands in north Sulawesi, apparently
naturally, albeit from mainland populations (Helgen and
Jackson 2015), and in the Sula islands S. pelengensis is recorded
from open dry forest, secondary habitats, farmland and scrub,
DAustralian Mammalogy T. E. Martin et al.
matching the habitats that dominate Manui (Flannery 1995;
Helgen and Jackson 2015).
Not only do south-east Sulawesis offshore islands support
populations of endemic cuscuses, but they may also represent
particularly important conservation areas for these species.
Buton, in particular, still supports nearly 350 000 ha of lowland
tropical forest, 65% of which lies within ofcial protected areas
and much of which is considered to have high conservation
value (Cannon et al.2007; Martin et al.2015). Therefore, on
Buton at least, large tracts of suitable habitat remain for both
A. ursinus and S. celebensis an important refuge given rapid
habitat loss elsewhere in their range (Helgen et al.2008; Salas
et al.2008; Helgen and Jackson 2015).
Additionally, our observations also indicate that all three
cuscus species recorded are capable of utilising, at least to
a certain degree, degraded forest, scrub, farmland and garden
habitats, in some cases where these non-forest habitats have
little or no connectivity to closed canopy forest ecosystems.
Such utilisation has been found for other Sulawesi endemics
(Martin and Blackburn 2010; Gillespie et al.2015), and has
been previously described in S. pelengensis (Helgen and
Jackson 2015) as well as in other cuscus species in New Guinea
(e.g. Leary et al.2016b). However, these patterns have not
been previously reported in A. ursinus and S. celebensis, with
both Salas et al.(2008) and Helgen and Jackson (2015) stating
that these species do not readily use disturbed habitats. Our
results raise the possibility that these species may be more
adaptable to habitat modication than previously thought,
although how optimal these habitats are for these cuscus species,
and the extent to which they are used, remains unclear.
The other important threat to these species hunting for food
(Helgen et al.2008; Salas et al.2008; Helgen and Jackson 2015)
may be a lesser concern in south-east Sulawesi compared with
elsewhere. MacKinnon (1979), OBrien and Kinnaird (1996),
Riley (2002) and Lee et al.(2005) all report that both A. ursinus
and S. celebensis are heavily hunted for food in predominantly
Christian northern Sulawesi. On south-east Sulawesis offshore
islands, however, where populations are predominantly Muslim,
hunting pressures may be lower, as consuming cuscus meat is
considered a religious taboo (Lee et al.2005). The minority non-
Muslim communities also do not appear to eat cuscus meat on
Buton, despite not being explicitly constrained by religious
beliefs (Priston 2005). Support for the suggestion that hunting
pressure is lower is indirectly provided by the fact that, in the
course of our cumulative 13 eld seasons in the region, we have
never observed cuscus being actively hunted, or sold in a market,
as a source of food on any of our study islands. Community
interview data presented in previous research (Priston 2005;
Hardwick et al.2017) also indicate that, unlike pigs and
macaques, farmers on Buton do not consider cuscuses to be
signicant crop raiders, and they are not persecuted. Care should,
however, be taken when interpreting the results of these
interviews, given that participants may have provided responses
they believe to be appropriate when discussing conservation
issues, rather than reporting the truth (Hardwick et al.2017).
Nevertheless, people on Buton have frequently told us in
conversation that they have hunted or otherwise persecuted
several threatened and protected species on the island, including
the lowland anoa (B ubalus depressicornis), maleo (Macrocephalon
maleo) and booted macaque (Priston 2005; DGT and PMW,
pers. obs.) but never any cuscus species. Rare second-hand
accounts exist of A. ursinus being captured and kept as pets
on Buton and Kabaena (although none of the authors have
personally observed this). It is perhaps possible that a hypothetical
lower hunting pressure on cuscus in our study area may also help
explain their presence in a range of non-forest habitats here,
anding that has not been reported elsewhere. It could be that in
northern Sulawesi (where most previous eld data for these
species have been sourced), cuscus are principally found in forest
habitats as hunting has extirpated them from other, more open,
habitats where they are more conspicuous and easily harvested,
not because they are intrinsically ecologically dependent on
these forest habitats. However, although we have never observed
evidence of hunting, we cannot be sure this never occurs
without completing targeted surveys, as conducted by Lee et al.
(2005). All cuscus species in south-east Sulawesi breed
slowly, typically raising a single offspring at a time, which
takes several years to reach maturity (Helgen and Jackson 2015).
Thus, even a low or moderate level of hunting pressure that
remained undetected during our eldwork might still exert
a substantial inuence on their demography.
In summary, our ndings suggest that the islands of south-east
Sulawesi are likely to represent important strongholds for at
least two of the regions endemic cuscuses, supporting large
areas of habitat and potentially experiencing lesser hunting
pressures than reported elsewhere in these speciesranges.
Conicts of interest
The authors declare no conicts of interest.
Acknowledgements
This project was supported by Operation Wallacea. We thank the Indonesian
Institute of Sciences (LIPI) and Kementerian Riset dan Teknologi Republik
Indonesia (RISTEK) for providing a series of permits for 20042014
eldwork on Buton, and permits 174/SIP/FRP/E5/Dit.KI/V/2016 and 160/
SIP/FRP/E5/Fit.KIVII/2017 for all other islands. We also thank David Kelly,
Nicola Marples, Vivien Cumming, and two anonymous reviewers for their
constructive and helpful comments.
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FAustralian Mammalogy T. E. Martin et al.
www.publish.csiro.au/journals/am
... It is the most primitive and largest species within the Phalangeridae family (Shepherd et al. 2018). The Sulawesi bear cuscus is endemic to Sulawesi and several satellite islands (Martin et al. 2019), where it inhabits undisturbed lowland tropical rainforest and (less frequently) plantations and gardens (Helgen & Jackson 2015;Martin et al. 2019). ...
... It is the most primitive and largest species within the Phalangeridae family (Shepherd et al. 2018). The Sulawesi bear cuscus is endemic to Sulawesi and several satellite islands (Martin et al. 2019), where it inhabits undisturbed lowland tropical rainforest and (less frequently) plantations and gardens (Helgen & Jackson 2015;Martin et al. 2019). ...
... Ailurops ursinus is listed as Vulnerable (Salas et al. 2019;Suyanto et al. 2020), being threatened with extinction from illegal wildlife trade, hunting and deforestation, with continuing decline of populations (Saragih et al. 2010;Dahlen 2013;Shepherd et al. 2018;Martin et al. 2019;Salas et al. 2019). The large areas of habitat available and reduced hunting pressures on islands in southeast Sulawesi are likely to be the strongholds for A. ursinus; for example, A. ursinus on Buton island in southeast Sulawesi are widespread and common (Martin et al. 2019). ...
Article
en Possum acoustic behaviour is complex and varies substantially, with some species having numerous calls used in various contexts, while other species are limited to one known vocalisation or non‐vocal sounds. Here, we report that the first known recording of Sulawesi bear cuscus (Ailurops ursinus, 1824) acoustic behaviour and observations of associated behavioural displays. We observed an animal adopting a raised posture in apparent threat display and audio‐recorded the concurrent short, harsh sounds. The animal produced a ‘chatter’, composed of four notes given at short, regular intervals, followed by a series of ‘clicks’ given at longer and irregular intervals. We describe the frequency and temporal characteristics of these sounds. Clicks were variable in acoustic structure, possibly falling into three subtypes, and some clicks overlapped in acoustic features with individual chatter notes. Click and chatter notes were broadband and non‐tonal, and so appear to be non‐vocal sounds, produced by the mouth or tongue rather than larynx. Our observations and recording of A. ursinus contribute to the natural history of this poorly known and enigmatic species, that is currently threatened with extinction from illegal wildlife trade, hunting and deforestation. Abstrak id Perilaku akustik possum bersifat kompleks dan sangat bervariasi, dengan beberapa spesies memiliki beragam panggilan yang digunakan dalam berbagai konteks, sementara spesies lain diketauhi terbatas pada satu suara vokalisasi atau non‐vokal. Di sini, kami melaporkan rekaman pertama yang diketahui dari perilaku akustik Kuskus beruang Sulawesi (Ailurops ursinus,1824) dan pengamatan dari tampilan perilaku terkait. Kami mengamati seekor hewan yang mengadopsi postur tubuh yang tampak seperti ancaman, dan merekam audio suara yang pendek dan kasar secara bersamaan. Hewan itu mengeluarkan "obrolan", terdiri dari empat nada yang diberikan secara singkat dan teratur, diikuti oleh serangkaian suara decak yang diberikan dengan interval yang lebih panjang dan tidak teratur. Kami menjelaskan frekuensi dan karakteristik temporal suara‐suara ini. Suara decak bervariasi dalam struktur akustik, mungkin terbagi dalam tiga sub‐jenis, dan beberapa suara decak tumpang tindih dalam fitur akustik dengan catatan obrolan individual. Nada decak dan obrolan adalah nada pita lebar dan non‐nada, sehingga tampak seperti suara non‐vokal, yang dihasilkan oleh mulut atau lidah daripada laring. Pengamatan dan pencatatan kami terhadap A. ursinus berkontribusi pada sejarah alami spesies yang kurang dikenal dan misterius ini, yang saat ini terancam punah karena perdagangan illegal satwa liar, perburuan dan penggundulan hutan. Abstract in Indonesian is available with online material.
... Keterangan ( Kepadatan populasi dapat dihitung menggunakan rumus (Southwood & Henderson, 2000): Boonratana (1999), bahwa distribusi kelompok primata pemakan daun dipengaruhi oleh ukuran kelompok dan ketersediaan pakan. Indikator habitat yang baik adalah habitat yang mampu menyediakan sumber pakan yang cukup dari segi kelimpahan jenis vegetasinya serta jumlahnya (Martin et al., 2019;Kiroh, Hendrik, Ratulangi, & Rimbing, 2021). Daya dukung habitat berupa produktivitas sumber pakan sebagai penyokong terhadap satwa yang tinggal pada suatu habitat (Kuswanda & Bismark, 2007;Basalamah et al., 2010). ...
Research
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Silvery lutung (Trachypithecus cristatus) is one of the primates of Cercopithecidae family distributed in Kalimantan and Sumatra. Its habitats are coastal areas, mangroves, shorelines and riparian areas. Currently, research on the silvery lutung has been widely carried out on Sumatra island, but there is a lack of information about the species on Borneo Island. One of the interesting habitats to study is the restored mangrove forest. This study aimed to determine the distribution and population density of the silvery lutung and identify the type of feed in Handil Baru Village, Samboja Sub District, Kutai Kertanegara District, East Kalimantan. Initial information that can be extracted is in the form of an ecological study. The population density was determined by direct observation using the Concentration Count method, while the distribution was analyzed using ArcGIS software version 10.4. We found three groups of silvery lutung in the mangrove area of Handil Baru Village, scattered in Handil Baru Muara, Raden River, and Mantri River. The average population density in each area was 6.63 individuals/ha; 2.65 individuals/ha and 0.72 individuals/ha, respectively, with the number of individuals per group ranging from 17-24. There were 22 plant species found as food sources for the lutungs. The population density was influenced by the extent of habitat use, the number of individuals at the location, and food availability. This research can be used as a reference for habitat management and regional development that contributes to the preservation of silvery lutung.
... Previous study has also found cuscus that mostly occupied the forest areas that have been converted into plantations in Obi Island because the availability of feed sources (Tamalene et al., 2019). The factors of feed availability in habitat indicators are the abundance of species and numbers (Martin et al., 2018), as well as the flowering season in October-December (Marthinus & Tuaputty, 2015), while the time of our study carried out in November-January with the dominance of fruit of the fig species (Ficus spp.). The results of the maximum likelihood classification showed that the area of each habitat suitability class in each logged block did not show a graph of decrease or increase based on the year of logging. ...
Article
Previous studies are most focusing on the species identification and habitat used by cuscus, while habitat changes are the most threats on cuscus. Habitat changes caused by the operation of logging concession such as PT Wijaya Sentosa creates an impact to the habitat selection of cuscus, which is highly dependent on forest cover. Efforts to protect cuscus species in production forests need to be carried out to ensure their sustainability habitat. This study aims to determine habitat preference of cuscus and model the suitability habitat of cuscus. The Neu index calculation is used to measure the level of habitat preference, while the habitat sustainability map is modelled by biotic, abiotic, and human disturbance components into MaxEnt application. Vegetation data is analyzed to describe tree density, tree richness, and strata using of cuscus by SexFI application. The study showed that the most preferred habitat by cuscus is LoA 2018 because LoA 2018 had the most species richness so cuscus might be able to choose their vegetation food. The habitat suitability of cuscus in PT Wijaya Sentosa covers 21,116.59 ha and didn’t follow the pattern of increasing Et+ logging but was strongly influenced by logged blocks as much as 55.2% contribution.
... The only terrestrial vertebrate recorded was the invertivore reptile (Reptilia) Emoia atrocostata. Future research could include camera traps placed in the canopy to record the presence of important vertebrate herbivores, such as the cuscus family (Phalangeridae), although they are thought to be increasingly uncommon in the region due to hunting pressure (summarized in Martin et al. 2019). In addition, passive acoustic monitoring could be utilized to survey important top predators in this system, particularly the mangrove avifauna (Acevedo & Villanueva-Rivera 2006;Buelow & Sheaves 2015), allowing for the building of multi-level interaction network including predatory interactions, giving even greater insight into the structure and function of these mangrove systems. ...
Article
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Mangroves are uniquely important ecosystems, for preserving biodiversity, sustaining livelihoods and mitigating against climate change. However they are degraded globally and are therefore a priority for ecosystem restoration. To date, the assessment of mangrove restoration outcomes is generally poor, and the limited studies that do exist are focussed largely on forest area. Thus, more holistic ways of assessing the outcomes of mangrove restoration projects on biodiversity and associated ecological processes are urgently needed. Ecological networks are a useful tool for simultaneously examining both. Here, we assessed the utility of using species-interaction networks for evaluating mangrove restoration outcomes for the first time. We compared the structure and complexity of mangrove ecological networks in replicated ‘Monoculture Reforestation’, ‘Mixed Species Regeneration’ and ‘Reference Forest’ plots in two study areas in Sulawesi, Indonesia, an estuarine and a coastal fringe mangrove system. We also combined and evaluated sampling methods, utilising traditional plant-animal sampling while also integrating video recording data in a novel way. We found significant differences in the structure and complexity of mangrove networks between restored and natural plots, with contrasting effects between the two sites. Our results show differences in the complex ways in which taxa interact in mangrove restoration projects, which would be overlooked if common biodiversity metrics such as species-richness were used alone, with consequences for the restoration of ecosystem functioning. We also highlight the utility of video recording data collection for constructing species interaction networks, overcoming the detrimental impacts of observer presence for some key species. This article is protected by copyright. All rights reserved.
... The Zoological Society of London (ZSL) included this species as one of Evolutionarily Distinct and Globally Endangered species (EDGE) based on evolution, life history, and threat status (EDGE 2018). Therefore, this endemic cuscus represents an important global conservation priority (Martin et al. 2018). Within Indonesia, A. melanotis is protected according to the Republic of Indonesia Minister of Environment and Forestry Regulations. ...
Article
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The Talaud bear cuscus (Ailurops melanotis) has been reported from Sangihe (the largest island in the Sangihe Island group) and Salibabu (within the Talaud Islands). As an endemic species of Indonesia, this species is rare and there is no certainty regarding its precise geographic distribution or population size. This research aimed to estimate population density and provide the first preliminary data on its geographical distribution, as well as general description of its habitat. Our research shows that A. melanotis occurs on three islands: Salibabu Island, Nusa Island, and Bukide Island, and probably also exists in the Sahandaruman mountain on Sangihe Island. Our population surveys estimate, population density on each island as: Salibabu: 3.69 ± 2.54 ind/km 2 , with an estimated total population of 28.95 individuals, Nusa Island: was 12.31 ± 2.58 ind/km 2 , with an estimated population of 19.08 individuals, and Bukide Island: 7.17 ± 1.79/km 2 , with an estimated population of 10.40 individuals. Information regarding population is a key guiding factor in conservation efforts, where population size is related to extinction risk (threat status) and its geographical distribution, this can help to determine conservation priorities for species or habitats.
... The forested core of Wawonii may be of high conservation value (Cannon et al. 2007) and the island has been shown to be important for other threatened taxa (Farida & Dahrudin 2008). Nearby Buton Island is home to some of the most biodiverse remaining lowland forest in Sulawesi, but is experiencing serious deforestation (Howard & Gillespie 2007, Martin et al. 2012, Martin et al. 2019. Much like Buton, Wawonii is experiencing continued habitat loss and degradation, lending urgency to the need for more rigorous surveying and protection of its almost uncharted biodiversity. ...
Article
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Despite being an important centre of endemism, the southeast peninsula of Sulawesi and its satellite islands have remained ornithologically neglected. While relatively extensive surveys have been carried out on Buton Island (the largest satellite of Southeast Sulawesi), the avifauna of much of the rest of the region is poorly understood. We visited the islands of Muna and Wawonii, and Lasada village on mainland Sulawesi, in the summer of 2017 to collect data for avian biogeographic research. The 2017 expedition combined transect surveys and mist-netting, allowing for a wide assessment of the avifauna at these sites. During these visits all bird species encountered were recorded, providing the first scientific assessment of the avifauna of Wawonii and providing much needed information on the avifauna of Muna Island and the southeast peninsula of Sulawesi. In total 119 species were recorded, of which 33 are regional endemics, two are classified as Near Threatened, two as Vulnerable and one as Endangered.
... Kondsi habitat yang baik akan memberikan dampak positif bagi kehidupan kuskus, begitu juga sebaliknya kondisi habitat yang tidak baik akan memberikan dampak negatif bagi kehidupan Kuskus. Indikator dari habitat satwa yang baik adalah tersedianya sumber pakan yang cukup, baik dari segi kelimpahan jenis maupun jumlahnya (Martin et al., 2018). ...
... Hasil penelitian menunjukan bahwa terdapat 39 individu Kuskus jenis Phalanger ornatus (Tabel 1,2,4,5,6,7,8) (Martin et al., 2018). ...
Article
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Indonesia has a very high fauna diversity. Cuscus is an Australian marsupial of the Phalangeridae family that has a quite extensive distribution in Indonesia particularly in the eastern part of the region. Its natural distribution is also found in Papua New Guinea to Australia. There are two genera disvovered in North Maluku, namely the genus Phalanger and Spilocuscus. Those genera are found in Halmahera Island, Bacan Island, Obi Island and Morotai Island. Cuscus in Obi Island is P. rothschildi that is distributed in the three islands: Obi, Bisa, and Obi Latu. This cuscus species is discovered in area with altitude of 100 m above the sea level. The study aimed to o analyze the population density of the Phalanger genus cuscus and identify its feed plants in Obi Island. The research was conducted using transect survey method and data collection techniques including observation, interviews and documentation. The results showed that cuscus in Obi Island was identified as Phalanger rothschildi (endemic species) and Phalanger ornatus. The cuscus density in the forest area of Jikotamo village was about 25 individual/ha. 16 plant species were eaten by the cuscus as the main food source. The most preferred plants were sirih hutan (Piper aduncum L.), kersen (Muntingia calabura L.) and Awar-awar (Ficus septica).
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The population of the bear cuscus Ailurops ursinus (Temminck, 1824), an arboreal marsupial endemic to Sulawesi, Indonesia, and its satellite islands, is declining rapidly due to poaching and habitat loss, even in protected areas. However, despite concerns over its persistence, little is actually known of this secretive species. This research investigated the characteristics of the selected habitats and diet of the bear cuscus in four ecosystem types (lowland non-dipterocarp forest, lowland limestone forest, lowland monsoon deciduous forest, lowland monsoon evergreen forest). Habitat use data were collected through direct encounters and indirect observations (tracks, signs, secondary information), and analyzed using a chi-square goodness-of-fit test. Habitat characteristics and diet availability were determined using vegetation analysis. Diet data were obtained using direct observations, feed remains, and interviews. The lowland non-dipterocarp forest ecosystem was used significantly more by bear cuscus populations. Its habitats across the four ecosystem types had similar environmental conditions. Fifty-five plant species, eaten mostly as young leaves and leaf buds, encompassed the bear cuscus’ diet, with the Moraceae family being the most representative. Considering the ubiquity of Moraceae in the bear cuscus’ habitat, these results highlight the impact unchecked deforestation will continue to have on lowland Indonesia and its endemic species.
Article
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Peninsular Malaysia is currently thought to host the highest biodiversity of Old World bats of any region, with 110 species recorded. However, the availability of literature to facilitate a similarly thorough species ‘checklist’ is not as readily available for other parts of Southeast Asia, including Sulawesi, Indonesia. Here we highlight 13 new species records from the long-term bat monitoring programme on Buton Island, South East Sulawesi, expanding on Patterson et al.’s (2017) previous inventory for this study area. One species (Hipposideros galeritus) is a new record for Sulawesi, and seven species (Cynopterus c.f. minutus, Rousettus celebensis, Megaderma spasma, Hipposideros c.f. ater, Myotis c.f. horsfieldii, Myotis c.f. moluccarum, and Myotis c.f. muricola) are new records for Buton Island. The remaining five species (Thoopterus nigrescens, Dobsonia exoleta, Acerodon celebensis, Mosia nigrescens, and Mops sarasinorum) have been previously reported from Buton but were missing from the prior site inventory. We also correct a probable mistaken species identification in the previous inventory (Cynopterus cf. titthaecheilus, now identified as Thoopterus nigrescens). This brings the total of confirmed species detected on Buton to 35, equating to 46.7% of all Sulawesi’s known bat diversity in c. 3% of its land area. We highlight Buton as a key area for conserving the region’s bat species.
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Human–wildlife overlap is increasing worldwide as a result of agricultural expansion. This can reduce human tolerance of wildlife, especially if wildlife threatens human food sources. The greatest threat to the declining populations of the endemic Buton macaque (Macaca ochreata brunnescens) is habitat destruction, but as a common crop-feeding species, there is also an additional risk of retaliation killings from farmers. Finding means of reducing this risk will thus help secure the long-term future of this range-restricted subspecies. Here, we investigate variability in farmers’ perceptions of primate crop-feeding and mitigation techniques in three farming communities on Buton Island, Indonesia, which differ in wealth and agricultural resources. We employ a mixed methodology, collecting qualitative social data from focus groups and quantitative observational data to measure macaque crop-feeding occurrences. Our findings indicate that the least wealthy community used lethal control methods more frequently than the comparatively wealthier communities, even when the crop-feeding problem was less severe. The least wealthy community also expressed high levels of fear of macaques, and had the most negative perceptions of them. This community also had no knowledge of the macaques’ conservation status or their ecological roles. We recommend that efforts to protect Buton macaques focus on education and the use of effective nonlethal mitigation techniques, such as electric fencing. We also suggest that initiatives to support such measures may be most effectively directed toward communities with relatively low economic wealth and high reliance on subsistence agriculture, especially where crop-feeding wildlife is feared, even when such communities do not experience the highest losses from crop-feeding wildlife.
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We studied the daily time budget and feeding activity of the bear cuscus, Ailurops ursinus, in the Tangkoko-Duasudara Nature Reserve, North Sulawesi, Indonesia. Bear cuscuses spent 63.4% of their time resting, and feeding accounted for only 5.6% of their activity. Bear cuscuses fed on 31 species of plants, including 26 identified trees and lianas from 17 families and 5 unidentified mistletoes. Dietary preference was influenced by availability of young leaves, and bear cuscuses maximized the amount of young leaves in the diet.
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Although commonly reservoired within Tertiary rocks, hydrocarbons in eastern Indonesia have generally been sourced from Mesozoic sediments deposited on the continental margin of northern Australasia. Fragments of this margin are now widely dispersed as allochthonous terranes throughout the area, one of the most far traveled examples being the island of Buton, southeast of Sulawesi. Asphalt reserves on Buton support a significant local industry and exploration continues for oil and natural gas. In common with other prospective Australian-derived terranes around the margins of the Banda Sea, Buton is now separated from Australia by the active plate boundary marked by the Java Trench and the collision trace along the Timor, Tanimbar, and Seram troughs. Buton differs from these other terranes in its distance from that boundary and its consequent insulation from the effects of the Pliocene-Pleistocene collision between Australia and the Banda arcs. Reconstruction of the geological history of the Buton terrane thus has an important role in guiding future exploration in the other Australasian fragments in eastern Indonesia. Geophysical studies of the Buton region have used seismic reflection, gravity, and magnetic (including paleomagnetic) techniques. Seismic reflection images generally record extension rather than compression as dominating the recent history of the area. Gravity data define the present-day western limits of the Buton terrane and suggest that in the east the terrane includes the almost entirely submerged Tukang Besi platform. The gravity surveys also demonstrate that the ophiolitic rocks exposed on Buton are not attached to deep roots, but are thin and isolated overthrust sheets. Therefore they do not mark a terrane boundary and their presence has little bearing on the prospectivity of the area. Paleomagnetic results document the independent movements of thrust sheets on Buton during the Pliocene-Pleistocene. The combined data from Buton record its separation from Australia as part of a microcontinental block in the Jurassic or Late Triassic, followed by collision with the Eurasian margin in southeastern Sulawesi in the Oligocene or early Miocene. Collision was followed by extension (as in Sulawesi itself) producing minor separation of Tukang Besi from Buton and much greater dispersion of other fragments of the microcontinent, some of which have since been incorporated in the new collision zone in the Outer Banda arc. The oil seeps and asphalt deposits of Buton are proof that hydrocarbons in the Banda arc fragments can be sourced from within these fragments and are not necessarily derived from the underthrusting Australian margin.
Article
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The issues of habitat loss and hunting are of paramount importance to wildlife conservation in Asia. In Sulawesi, Indonesia, these problems are having a serious impact on the vertebrate fauna. Using line-transect methods, the densities of 11 species of large birds and mammals were compared between 1979 and 1994 in the Tangkoko-DuaSudara Nature Reserve in North Sulawesi. During those 15 years, populations ofanoa Bubalus depressicornis, bear cuscus Phalanger ursinus, crested black macaque Macaca nigra, maleo Macrocephalon maleo and red junglefowl Gallus gallus declined by 50–95 per cent while populations of Sulawesi pig Sus celebensis, Tabon scrubfowl Megapodius cumingii, Sulawesi tarictic hornbill Penelopides exarhatus and red-knobbed hornbill Aceros cassidix increased by 5–100 per cent. We considered hypotheses for these changes: habitat loss outside the reserve, habitat degradation inside the reserve, and hunting. Only hunting adequately explained the pattern of changing densities observed. Unless protection from hunting is enforced for these species, we may soon witness the demise of these unique animals in North Sulawesi and possibly throughout the island.
Article
Surveys between 1995 and 1999 brought the number of mammal species known to occur on the remote Sangihe and Talaud islands, Indonesia, from 34 to 37, of which 30 are indigenous and 22 are bats. Populations of bear cuscus Ailurops ursinus and Sulawesi small cuscus Strigocuscus celebensis are represented by endemic subspecies, whilst five little-studied species (Talaud Islands flying fox Acerodon humilis, Sangihe tarsier Tarsius sangirensis, Sangihe squirrel Prosciurillus rosenbergi, short-tailed Talaud melomys Melomys caurinus and long-tailed Talaud melomys M. talaudium) are endemic to the archipelago. Two squirrel species were recorded on Sangihe for the first time: Sulawesi dwarf squirrel P. murinus and Sulawesi giant squirrel Rubrisciurus rubriventer. In total, eight species occurring on the islands are categorized as globally threatened on the 2000 IUCN Red List of Threatened Species. Habitat loss and hunting are the main threats on the Sangihe islands, where only 800 ha of primary forest remain. Large areas of Karakelang, in the Talaud Islands, are still forested, and a 24,669 ha wildlife reserve has been recently established. The main pressure facing mammal species on the Talaud Islands is hunting, particularly mist-netting of fruit bats for local consumption and trade. In order to control hunting and prevent further forest loss and degradation, future conservation efforts should focus on community-based conservation, in particular raising community awareness and increasing law enforcement. Two concurrent projects are now tackling some of these issues.
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The wildlife of the island of Sulawesi in Indonesia is a remarkable mixture of Asian and Australasian forms, with many endemic species. But it is being rapidly destroyed as a result of timber logging, hunting and agriculture. However, Dr MacKinnon, who is manager for two World Wildlife Fund projects in North Sulawesi, believes that action now could save most of the larger animals. With Indonesia's new interest in conservation he is optimistic that this will be achieved.
Article
We examined the distribution and broad habitat associations of the herpetofauna on three offshore islands of southeast Sulawesi, Indonesia. A total of 74 amphibian and reptile taxa were recorded, comprising 13 frogs, 29 lizards, 29 snakes, 1 freshwater turtle, and 1 crocodile. Of the total taxa, 38 percent were endemic to Sulawesi, 13 were new undescribed taxa. Range extensions were also recorded for one taxon previously not known from Sulawesi. Herpetofauna of these islands is largely derived from that of mainland Sulawesi, and as for Sulawesi generally, is depauperate compared with herpetofaunal assemblages in Borneo, Java, and Thailand. Taxon richness was much higher in minimally disturbed forest and forest habitats with only moderate disturbance levels than in highly modified or disturbed habitats, such as secondary forests, plantations, and villages. Disturbed habitats were characterized by widespread, habitat generalists and human commensals. Forests were characterized by endemic and habitat-specialist taxa. Little discrimination of taxon composition or endemism was found between minimally and moderately disturbed forest habitats. These results reaffirm the need for more general biological survey and research in this region. Taxa most likely to be displaced by human impacts tend to be endemic taxa, for which there exists little or no ecological information. The similarity in the herpetofaunal community structure between habitats with minimal and moderate disturbance levels has important implications for our understanding of ecological resilience in tropical herpetofaunal communities.