Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 May 2021 | 13(6): 18419–18426
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.6271.13.6.18419-18426
#6271 | Received 05 June 2020 | Final
received 06 April 2021 | Finally accepted 15 May 2021
Status of Sumatran Tiger in the Berbak-Sembilang landscape (2020)
Tomi Ariyanto
1, Yoan Dinata 2,
Dwiyanto 3, Erwan
Turyanto 4, Waluyo
Sugito 5, Sophie Kirklin
6 & Rajan Amin 7
1–3, 5–7 Zoological Society of London,
Outer Circle, Regent’s Park, London, NW1 4RY.
4 Berbak Sembilang
National Park Agency, Jambi, Indonesia.
1 tom.ariyanto@gmail.com
(corresponding author), 2 yoan7dinata@gmail.com, 3 dwiyantoduwez@gmail.com,
4 waluyosugito82@gmail.com, 5 tourerwan@gmail.com, 6
skirklin@icloud.com, 7 raj.amin@zsl.org
Editor: L.A.K. Singh, Bhubaneswar,
Odisha, India. Date of publication: 26 May
2021 (online & print)
Citation: Ariyanto,
T., Y. Dinata, Dwiyanto, E.
Turyanto, W. Sugito, S. Kirklin & R. Amin (2021). Status of Sumatran Tiger in the Berbak-Sembilang landscape (2020). Journal of Threatened Taxa 13(6): 18419–18426. https://doi.org/10.11609/jott.6271.13.6.18419-18426
Copyright: © Ariyanto
et al. 2021. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: Panthera; GEF-UNDP Tiger Project; IUCN-KfW ITHCP Project; Disney Conservation
Fund.
Competing interests: The authors
declare no competing interests.
Author details: Tomi Ariyanto’s
educational background is in biology with an emphasis on conservation and
ecology. He holds an MSc from the University of Indonesia. Tomi has 13 years of
conservation experience, largely in Sumaran tiger and
Bornean orangutan conservation. From 2014-2020, he joined the tiger
conservation project ZSL Indonesia Program, responsible for the implementation
of tiger population monitoring and area protection. Now, Tomi is a government engagement officer
at Borneo Nature Foudation. Yoan Dinata holds an MSc in
Conservation Biology from DICE, Kent University. 14 years of conservation
experience focuses on large mammal conservation including Sumatran tigers,
elephants and tapirs. Since 2008 he has been an active member of the Sumatran Tiger
Conservation Forum “HarimauKita”. He is also a former
chairman for the forum. His fields of
interests include felid population ecology, habitat management and wildlife
trade. From 2014-2020, Nata was the Tiger Conservation Project Manager in ZSL Indonesia
Program. Dwiyanto
holds BSc from Andalas
University. From 2016-2020 he was a field team member with the ZSL Indonesia
Program, using his expertise and experience in field survey and data
management. Dwi is now a camera trapping leader at
Fauna & Flora International Riau Program. Waluyo Sugito is a member of the
community surrounding BSNP. He worked for ZSL Indonesia Programme
from 2010-2020, starting in 2010 as field staff and progressing to Field team
leader and GIS / Remote Sensing specialist in the ZSL Indonesia Tiger
Conservation Program. Erwan Turyanto is
the Forest Ecosystem Technician Coordinator of Berbak-Sembilang
National Park Agency. He has 19 years of experience in area management and
wildlife conservation in BSNP. Erwan holds magister
of Agri-business from the University of Jambi.
Sophie Kirklin
has a masters in Human Evolution and Behavior from
University College London. She worked for ZSL supporting projects in Indonesia
for over 2 years, and previously coordinated an orangutan behavioural
research project in Central Kalimantan for 1.5 years with the Borneo Nature
Foundation. Sophie is now a Research Associate at RBG Kew, delivering capacity
building courses to Indonesian partner institutes as part of project ‘Unlocking
the potential of Seasonal Forests to underpin Wallacea’s
green economy’. Rajan Amin is a senior wildlife biologist at
the Zoological Society of London with over 25 years of experience in African
and Asian grassland and forest ecosystems and in developing long-term conservation
projects for threatened species.
Author contributions: TA and YD conceived and designed
the study; TA, ET, D, WS assisted in data collection; RA and TA performed the
analysis; RA, TA and SK wrote the manuscript.
Acknowledgements: We thank the Indonesian Ministry
of Environment and Forestry, and Head of Berbak-Sembilang
National Park for granting permission to conduct the field surveys. We thank Edi Susanto, Zainal Abidin, Bagus Hutajulu,
M. Azwi Nardiansyah, Sismanto, Raziko, Suryadi & Saryono, and the
rest of the field team for assisting in the camera trap surveys. Our thanks also go to Citra Panjaitan, Iding Achmad Haidir and M. Anderi for their contribution to the Berbak
tiger monitoring programme. The
programme is supported by various donors including Darwin Initiative, Panthera, GEF-UNDP Tiger Project and IUCN-KfW.
Abstract: Monitoring the status of the
Critically Endangered Sumatran Tiger Panthera
tigris sumatrae is a
key component for assessing the effectiveness of conservation interventions,
and thus informing and adapting strategic planning for the remaining 600
Sumatran Tigers on the island. The Berbak-Sembilang National Park is an integral part of the
priority Berbak-Sembilang Tiger Conservation
Landscape, in a unique
habitat of mixed peat and freshwater swamp in eastern Sumatra. Our camera trap survey covered both the Berbak and Sembilang Tiger Core
Areas (BTCA, STCA) over a period of 10 years, with surveys undertaken in 2010,
2015, 2018–2019. The most recent
population density estimates (BTCA 1.33 adults/100 km2, 95% CI
0.82–1.91 with 19 adults; and STCA 0.56 adults/100 km2, 95% CI
0.45–0.89 with five adults) confirmed a small but stable population. A landscape level management approach is a
priority for tiger population recovery, consolidating ground-based protection
and establishing a well-maintained fire management system with reforestation of
affected areas along with multi-stakeholder engagement and partnerships. The study also recommends extending the BTCA
to include the primary swamp forest in the north of the national park, based on
evidence from camera trap surveys.
Keywords: Abundance, camera trap, density, Panthera tigris sumatrae, Sumatra.
Bahasa Indonesia: Harimau Sumatera telah
mengalami menurun populasi sekitar 16,6% antara tahun 2000 dan 2012 yang sebagian besar disebabkan oleh hilangnya hutan dan degradasi
habitat. Saat ini mereka sangat terancam
punah dan pemantauan yang efektif terhadap populasi tersisa dalam lanskap
prioritas sangat penting untuk menginformasikan
rencana pengelolaan dan konservasi guna mengamankan masa depan spesies karismatik
ini. Survei dengan kamera
penjebak di bentang alam prioritas konservasi harimau Berbak-Sembilang di Sumatera timur,
mencakup Area Inti Harimau Berbak dan Sembilang
(BTCA, STCA) selama 10 tahun.
Perkiraan kepadatan populasi 2018-2019 terbaru, yaitu di BTCA 1,33 indvidu dewasa / 100 km2 (95% CI 0,82 - 1,91) dan
STCA 0,56 dewasa / 100 km2 (95% CI 0,45 - 0,89) dengan perkiraan kelimpahan, BTCA 19 individu dewasa (95% CI 11 - 27) dan di
STCA 5 individu dewasa (95%
CI 4 - 8),hasil ini telah mengkonfirmasi bahwa populasi kecil tapi stabil.
Kepadatan populasi harimau Berbak-Sembilang lebih tinggi daripada
lanskap Kerinci-Seblat di
Sumatera Barat dan lanskap Lueser-Ulu Masen di Sumatera Utara, dan
lebih rendah dari lanskap Bukit Barisan
Selatan di Sumatera Selatan. Populasi harimau Berbak-Sembilang, seperti di lanskap konservasi harimau Sumatera lainnya, terancam oleh deforestasi, yang sebagian besar diakibatkan oleh kebakaran hutan secara rutin dan
pembukaan lahan untuk pertanian, dan perburuan ilegal
yang berkelanjutan, yang didorong
oleh perdagangan satwa liar ilegal nasional dan internasional.
Sejak tahun 2007, program konservasi harimau Berbak-Sembilang telah dilaksanakan melalui kerangka kerja sama antarlembaga. Pemantauan status populasi harimau merupakan komponen kunci untuk mengukur efektivitas intervensi konservasi di lanskap dan menginformasikan perencanaan strategis. Populasi harimau kecil lebih rentan
terhadap fluktuasi demografis dan genetik, yang dapat memengaruhi kelangsungan hidup jangka panjang
mereka. Oleh karena itu, pendekatan pengelolaan tingkat lanskap yang efektif menjadi prioritas, membangun koridor harimau yang berfungsi dengan lanskap harimau lainnya dan melindungi zona penyangga taman nasional. Sama pentingnya untuk meningkatkan kapasitas pemantauan, analisis data dan pelaporan di antara para pemangku kepentingan dan membangun basis data harimau tingkat nasional yang tersentralisasi untuk pemantauan status dan pengambilan keputusan.
Introduction
Among the six extant subspecies of tigers, the Sumatran Tiger Panthera tigris sumatrae survives in isolated populations across 27
forest patches in Sumatra (Figure 1, combined forest area ~140,226 km2)
(Wibisono & Pusparini
2010). It is listed as Critically
Endangered on the IUCN Red List (Linkie et al. 2008),
threatened by: 1) habitat loss and
fragmentation, largely the result of regular forest fires and land clearance
for agricultural use; 2) poaching, encouraged by the illegal wildlife
trade and international illegal wildlife
trade; and 3) human-tiger interaction (Nyhus et al. 2004; Ng & Nemora
2007; Linkie et al. 2008; Kartika 2017). To conserve the subspecies, six of the
existing 12 tiger conservation landscapes (TCLs) in Sumatra have been
designated as priority tiger landscapes in the National Tiger Recovery Plan,
including the Berbak-Sembilang TCL (GTI 2012), the
focus of the current study (forest patch number 26 in Figure 1).
The Berbak-Sembilang TCL comprises
Berbak-Sembilang National Park (BSNP, 3,442km2),
Air Hitam Peatland Protected Forest (187km2),
Orang Kayo Hitam Forest Park (181km2),
timber concessions (622km2), acacia plantation concessions (517km2),
and oil palm plantations (106km2) (KLHK 2020). With over 3,800km2 of wetland
forest (Wibisono & Pusparini
2010), the landscape is a combination of mixed peat swamp, freshwater forest (Giesen 2004; GTI 2012) and mangrove forest (Silvius et al.
2018a). It is also an important carbon
sink (GTI 2012), but it is experiencing deforestation caused by logging and
human-caused fires (Giesen 2004).
Since 2007, a tiger conservation programme has
been operational in Berbak-Sembilang TCL to monitor and protect the tiger
population and its habitat. The adaptive patrol management system, SMART
(Spatial Monitoring and Reporting Tool, https://smartconservationtools.org),
has been implemented by BSNP since 2014, led by the Tiger Patrol and Protection
Units (TPPUs), to tackle tiger poaching and habitat destruction. A Wildlife Conflict Response Team (WCRT) has
been operational since 2011 and a Wildlife Crime Investigation Unit (WCIU)
since 2015, combating the illegal wildlife trade, mitigating human-wildlife
conflict and combating the illegal wildlife trade. Establishing and conserving a viable tiger
population requires long term monitoring of tigers, co-predators, prey and
their habitats to evaluate a conservation programme’s
effectiveness and inform management decision-making (Jhala
et al. 2009; Goodrich et al. 2013). The
objective of the study was to assess the status of the Sumatran Tiger in the Berbak-Sembilang TCL, as part of implementation of a tiger
conservation strategy.
Materials and
methods
Study area
The study was conducted in Berbak Tiger Core
Area (BTCA, area 657km2) and Sembilang
Tiger Core Area (STCA, area 695km2) within BSNP, located on the east
coast of Sumatra island, Indonesia (1.08°–-2.45°S and 103.80°–-104.90°E). Berbak Tiger Core
Area mainly consists of freshwater swamp forest and peat swamp forest. The
topography is flat with elevation less than 15m (Giesen
2004). Sembilang
Tiger Core Area contains the largest area of mangrove forest in the
Indo-Malayan region (Silvius et al. 2018a).
It is made up of peatland and mangrove forest in the east and peat swamp
forest in the west. Both core areas come
together in the small blackwater Benuh River*, and in
a large peat dome in the west that forms the upper catchment of the Benuh River. Annual
rainfall is c. 2,466mm with lowest and highest records of 933mm and 3,972mm,
respectively (Silvius et al. 2018a).
Tiger
prey species in Berbak-Sembilang TCL
include Wild Boar Sus scrofa, Bearded
Pig Sus barbatus, Southern Red Muntjac Muntiacus muntjak,
Sambar Deer Rusa unicolor and two
sympatric species of chevrotain: Greater Oriental Chevrotain Tragulus napu and
Lesser Oriental Chevrotain Tragulus kanchil. The
other felid species in the study area include Sunda
Clouded Leopard Neofelis diardi, Marbled Cat Pardofelis
marmorata, Leopard Cat Prionailurus bengalensis, and Flat-headed Cat Prionailurus
planiceps (BSNP & ZSL 2018).
* A blackwater river is a river with a
slow-moving channel flowing through forested swamps or wetlands, whose water is
tannin-stained, in this case from the peat.
Field methods
Tiger population densities were estimated using
standardized camera trapping procedures based on capture-recapture method (Karanth
& Nichols 2002). Surveys were conducted in BTCA between 31
January 2018–22 August 2018, 14 February 2015–8 April 2016, and 23 June 2010–2
February 2011; and in STCA between 1 January–1 July 2019 and 5 September
2018–21 December 2018. A nine km2
grid system was used, and paired camera traps (1 pair of camera traps= 1 camera
trap station) were deployed in accessible tiger habitat (Figure 2). Average Sumatran Tiger home range is
estimated at 50–70 km2 for adult female and 110km2 for
adult male (Franklin et al. 1999).
Camera traps were placed along animal trails to maximise tiger detection
(Sunarto et al. 2013). Camera traps were attached to trees and
positioned approximately 40cm above ground with each of the paired camera traps
about 7m apart and pointing towards each other, in order to capture both flanks
of a tiger to facilitate individual identification. In total, 125 camera trap stations were
placed in BTCA and 146 camera trap stations in STCA over the five surveys
(Table 1). DLC Covert II, Panthera V4, and Reconyx Hyperfire HC500 digital cameras were used, programmed to
take three photographs per trigger with no delay. All camera traps used white flash to obtain
colour images at night to aid in individual tiger identification.
Data analysis
The metadata (i.e., image name, date and time)
associated with all images were extracted with Exiv2 software (Huggel 2012) and compiled in an Excel spreadsheet
(Microsoft Office Professional Plus 2010).
Information on identified species was then added for all images
obtained. Individual tigers were
identified based on their unique stripe patterns and gender determined by
secondary sexual traits.
Tiger population densities were estimated using
the Bayesian Spatial Capture Recapture (SCR) method. A 90-day data subset for each survey was used
for the analysis to avoid violation of the population closure assumption. Three data input files were created: a camera
trap station activity file specifying camera trap station location and operation
by 24-hour day; a tiger capture history file specifying capture events as a
single detection of an individual tiger at a camera trap station in each
24-hour period; and a binary habitat mask file. The analysis was carried out in
the program JAGS (Just Another Gibbs Sampler) accessed through the program R,
version 3.6.0 (R Development Core Team 2019) using the package RJAGS (http://mcmc-jags.sourceforge.net). In data augmentation, M was set to 80 –
larger than the largest possible population size (i.e., the number of activity
centres). The centroid of capture
locations of individual animals caught were used as the starting values for
activity centres. Three MCMC (Markov Chain
Monte Carlo) chains with 60,000 iterations, a burn-in of 1000, and a thinning
rate of 10 were implemented. This
combination of values ensured an adequate number of iterations to characterise
the posterior distributions. Chain
convergence was checked using the Gelman-Rubin statistic (Gelman et al. 2004),
R-hat, which compares between and within chain variation. R-hat values below 1.1 indicate convergence
(Gelman & Hill 2006). The approach
of Royle et al. (2014) was used for the model
goodness-of-fit test, calculating three statistics, all using Freeman-Tukey
discrepancies: individual animal by camera trap station capture frequencies,
aggregating the binary daily capture data by animals and camera trap stations;
individual animal capture frequencies, aggregating for each animal; camera trap
station animal capture frequencies, aggregating for each camera trap station.
For tiger prey species, trap rate for each species was calculated as the
mean number of independent photographic ‘events’ per trap day x 100. An ‘event’ was defined as any sequence for a
given species occurring after an interval of >60 min from the previous
three-image sequence of that species (Amin et al. 2015).
Results
Survey effort ranged from 1,152 trap nights
(BTCA 2010 survey with 27 camera trap stations) to 6,570 trap nights (STCA 2019
survey with 73 camera trap stations; Table 1).
The number of adult tigers captured at a study site ranged from 4 to 10
individuals (Table 1). One individual
was captured in both BTCA and STCA in different years.
Given the small sample size, it was not possible
to model differences in space use and movement range between the sexes. The STCA 2018 survey had only one individual
tiger captured at more than one location and therefore it was not
analysed. The Bayesian p0~1.sigma~1
model R-hat values for all estimated parameters were below 1.01 and fitted well
to the data (P=0.3-0.5 for all three statistics). Berbak Tiger Core
Area had a higher density of tigers (1.33 individuals/100km2, 95% CI
0.82–1.91 in 2018) than STCA (0.56 individuals/100km2, 95% CI
0.45–0.89 in 2019). The BTCA tiger population also showed a stable trend
between the study years (Table 2, 2.78% per year, SE 1.18).
The most recent BTCA (2018) and STCA (2019)
tiger population density maps, derived from the model, are not presented due to
data sensitivity2. Several
tigers were caught in 1–2 locations close to the northern periphery of BTCA,
and the western and southeastern edge of the STCA, so
there was greater uncertainty in their activity centres (Figure 3).
Our study provided evidence of
breeding, with two adult females each with two cubs photographed in STCA (2018–2019), and one adult female with her two
cubs in BTCA (2018) (Image 1). The
surveys also indicated relatively good population of medium-to-large-bodied
prey species, including Lesser Oriental Chevrotain, Wild
Pig, and Bearded Pig in BTCA and STCA (Table 3).
Discussion
Our study has shown that the tiger population
within BSNP is small, and has remained stable over the past ten years despite
facing significant threats (Giesen 2004). The estimated tiger density for BTCA was
similar to those recorded in the mangrove habitat of the Sundarban
landscape (1.08–4.79 tigers/100km2, Jhala
et al. 2016), while the estimated density of tigers in STCA was lower. Although we were unable to measure the scale
of hunting or poaching in BSNP, patrol data between 2015–2019 show that snares
were reported at a rate of 0.89 traps/100 km (BSNP 2020). A recent study on the spatio-temporal
distribution of human-tiger interaction in Sumatra classified the relative
distribution of conflict cases in the Berbak-Sembilang
TCL as low to moderate (Kartika 2017).
Furthermore, within the last five years, there has only been a single
record of human-caused tiger mortality in the landscape (Zoological Society of
London 2020). Evidence of tiger breeding
also suggests a relatively healthy prey population to support lactating
tigress.
Enhancing tiger
recovery in the Berbak-Sembilang landscape
Within the forests and peatlands of Berbak Sembilang TCL, habitat
loss, mainly by human-caused forest fires, is the current main threat (Abood et al. 2015).
In just eight years between 2000 and 2018, there were 12,084 fire
hotspots in BSNP (80% confidence level) occurring in both dry and wet seasons
(Mora et al. 2019). Across the
landscape, forest fires are changing the structure of the peat swamp forest
(Wetland International-Indonesia Programme 2002), with large areas of closed
canopy-tall trees with undergrowth being replaced by a mosaic of open patches
of grasses and shrubs (Giesen 2004). The average rate of annual forest loss for
the period 2010–2040 has been predicted to be 1.1–1.6 % (Elz
et al. 2015).
Maintaining forest integrity is critical for the
survival of tigers (Wibisono et al. 2011), and this
requires increased protection from illegal logging and forest clearance. A comprehensive fire management plan should
also be created and implemented, based on suitable technologies such as remote
sensing and appropriate levels of SMART patrolling in this physically
challenging environment. A reforestation
programme with replanting of indigenous trees needs to be urgently undertaken
in the affected areas. The BTCA should
be extended to include the primary swamp forest north of the area, following
camera trap surveys. The delineation of
the existing tiger core area was based on tiger occurrence detected by camera
trap and sign surveys (Wibisono et al. 2011; Goodrich
et al. 2013). A massive forest fire in
2015, however, has significantly altered forest cover in the core area, and our
study revealed activity centres of several tigers lying north of the core area
(Fig. 3).
Suitable habitat for tigers also needs to be
expanded, increasing the chances for establishing a long-term viable tiger
population. A landscape-based approach
to the management of the area is being implemented by the Indonesia Government,
combining the previously separate Berbak National
Park and Sembilang National Park into a single
national park (BSNP) under the management of a single authority (via MoEF decree No. P.07/2016).
The peatlands of Sembilang remain contiguous
with those of Berbak, and together they provide
habitat for tigers in this unique ecosystem (Silvius et al. 2018a). As a next step, it should be a priority that
government owned lands adjacent to the national park are protected, and
incompatible land-uses prohibited (Silvius et al. 2018b). Ultimately, concession areas will need to be
integrated into tiger managed habitats and habitat connectivity re-established
if tigers and other threatened wildlife are to have a future in this unique
landscape.
Table 1. Camera trap sampling
effort, number of tiger detections and number of individual adult tigers
captured (sub-period of 90 days was used for the analysis - see methods).
Area |
Survey year |
Trap-nights
(number of camera trap stations) |
Number of
tiger independent photo-captures |
Number of
adult tigers captured |
Number of
adult female tigers captured |
Number of
adult male tigers captured |
Berbak Tiger Core Area |
2018 |
2,885 (50) |
63 |
10 |
5 |
5 |
2015 |
3,731 (48) |
31 |
6 |
4 |
2 |
|
2010 |
1,152 (27) |
16 |
6 |
2 |
4 |
|
Sembilang Tiger Core Area |
2019 |
6,570 (73) |
11 |
4 |
2 |
2 |
2018 |
5,934 (73) |
13 |
4 |
1 |
3 |
Table 2. Estimates of Bayesian
spatial capture recapture model outputs.
Sigma is the ranging scale parameter.
Area |
Survey year |
Tiger
density (95% CI) per 100km2 |
Tiger adult
female density (95% CI) per 100km2 |
Tiger adult
male density (95% CI) per 100km2 |
Sigma (95%
CI) (km) |
Population
size (95% CI) |
Berbak Tiger Core Area |
2018 |
1.33 (0.82–1.91) |
0.66 (0.34–1.02) |
0.66 (0.34–1.02) |
4.16 (3.27–5.14) |
19 (11–27) |
2015 |
1.09 (0.48–1.78) |
0.70 (0.27–1.23) |
0.39 (0.14–0.82) |
3.02 (2.22–3.97) |
16 (7–26) |
|
2010 |
1.36 (0.54–2.35) |
0.49 (0.18–0.99) |
0.87 (0.36–1.63) |
3.82 (2.24–5.82) |
15 (6–26) |
|
Sembilang Tiger Core Area |
2019 |
0.56 (0.45–0.89) |
0.28 (0.22–0.45) |
0.28 (0.22–0.45) |
4.59 (2.67–6.97) |
5 (4–8) |
Table 3. Tiger prey species trap
rates for Berbak Tiger Conservation Area and Sembilang Tiger Conservation Area. The trap rate was calculated as the number of
independent photographic events per 100 trap days. We defined a camera trap ‘independent
photographic event’ as any sequence of photographs of the species occurring
after an interval of >60 minutes from the previous photograph of the species
(Amin et al. 2015).
Species |
BTCA 2010 |
BTCA 2015 |
BTCA 2018 |
STCA 2018 |
STCA 2019 |
Wild Boar Sus
scrofa |
0.66 |
0.79 |
1.81 |
6.07 |
5.80 |
Bearded Pig Sus
barbatus |
0.03 |
0.20 |
15.71 |
6.89 |
10.59 |
Sambar Deer Rusa
unicolor |
- |
- |
0.10 |
0.67 |
0.49 |
Muntjac Deer Muntiacus muntjak |
- |
- |
- |
0.16 |
0.01 |
Greater Oriental Chevrotain Tragulus napu |
4.33 |
0.35 |
9.81 |
0.74 |
0.35 |
Lesser Oriental Chevrotain Tragulus kanchil |
- |
0.19 |
13.62 |
5.56 |
4.17 |
Long-tailed Macaque Macaca fascicularis |
0.46 |
0.06 |
0.76 |
2.23 |
2.06 |
Pig-tailed Macaque Macaca nemestrina |
2.14 |
0.83 |
4.19 |
3.37 |
2.36 |
For
figures & image - - click here
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