See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/341380872

Dhole Cuon alpinus in Satchari National Park: on the first verifiable evidence
from northeast Bangladesh

Article  in  Mammalia · May 2020

DOI: 10.1515/mammalia-2019-0050

CITATIONS READS

2 536

3 authors:

Tania Zakir Harish Debbarma

University of Dhaka 3 PUBLICATIONS   2 CITATIONS   
10 PUBLICATIONS   3 CITATIONS   
SEE PROFILE
SEE PROFILE

Muntasir Akash
University of Dhaka
38 PUBLICATIONS   33 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Study of bird use of fig (Ficus species) in Satchari National Park View project

Assessment of bird diversity and their threats in the sandbars of Rajshahi metropolis View project

All content following this page was uploaded by Muntasir Akash on 08 September 2020.

The user has requested enhancement of the downloaded file.

Kalenga Wildlife Sanctuary of the Tarap Hill
TH

(Mukul et al. 2006). Having a tropical climate, the average
annual rainfall of the area is 4,162 mm. Temperature ranges
from about 12–32 °C, and relative humidity fluctuates between
74% during December to 90% during July–August (Choud-
hury et al. 2004; Mukul et al. 2006; NSP 2006).
The SNP is one of the smallest among the 17 national
parks of Bangladesh, an IUCN category II Protected Area
with an area undulating with slopes and hillocks, ranging
from 10–50 m in height (NSP 2006; WDPA 2016). The soil
texture is sandy loam to silty clay with sparse grass cover at
places. The entire park area is characteristically criss-
crossed by networks of small, dried out sandy-bedded
streams. These streambeds can hold temporary pools only
during the rainy season. Other than the monsoon, severe
water scarcity is a primary feature of the park and two
ponds excavated by the forest department serves the de-
mand of wildlife. The forest department has infrastructure
on the northwestern tip of the park. The only Tipperah
village of the area is also positioned there housing 24
families. The locals also face the issue of inadequacy of
water but never use the forest ponds. A national highway
traverses the reserve forest separating the northern limit of
the SNP from the rest. Five bridges were built through the
area to construct this highway. Dry streambeds run below
the bridge underpass, connecting the park with the RHRF.
The park is home to more than 208 plant species;
together with natural and mixed forests, the park is
bordered by tea estates, rubber and agar plantations
(Mukul et al. 2006). Southerly, the park shares a teak
plantation that merges with the borderline forests, then
with that of the Baramura Hill Range, Tripura (Figure 1A). A
total of six amphibians, 18 reptiles and 149 species birds
are deemed as denizens of the park. Of the 24 sighted
mammals of the park, 15 are carnivore including two ca-
nids, three herpestids, five viverrids, three mustelids, one
ursids, and three lesser cats (IUCN Bangladesh 2015; Khan
2018; NSP 2006). Till date, there has been no systematic
study on the biodiversity of the park.
Our findings of dhole came from our yearlong survey
(April 2018–April 2019) on the mammalian carnivores. We
deployed camera traps aided with sign survey and ques-
tionnaire survey. We used Bushnell 119576C Trophy Cam
HD Max Trail Cams and followed Wearn and Kapfer (2017);
Boitani and Powell (2012); O’Connell et al. (2011) and TEAM
Network (2011) for protocols regarding height and distance
among the camera trap stations. The maximum height of
the installed cameras was 30 cm above the ground. Each of
the cameras, corresponding SD cards, and locks were given
a unique identity (hereafter, deployment ID) to enable
correct notation of date, time and location of each of
photographic capture of a small carnivore.
MAMMALIA-2019-0050_proof  4 May 2020  9:24 pm
T. Zakir et al.: Dhole in northeast Bangladesh 3
We divided the study area into 53 grids of 250 m by
250 m and considered grids on the park border that
included more than 30% of the park area (Figure 1B).
Camera trap points were selected that we considered as
crucial after the initial reconnaissance survey. We
considered intersections of the trails include bridge un-
derpasses, trails, crossroads and wide sections of dried out
streambeds etc.
We carried out camera trapping by installing camera
traps in 12 grids for approximately 4 winter months
(October 2018–January 2019). We began with three camera
traps deployed from 25 October 2018 for more than a month
considering theft issues and, then, placed nine more
camera traps in different stations within the forest
(Figure 1B). Due to camera theft in three stations later on
and vandalization of one despite usages of python lock for
strengthened security, the total estimated sampling effort
turned out as 587 trap nights. We did not use any lure to
enhance the encounter rate. We sorted the pictures and
considered photos of 30 min or more apart as independent
events.
Considering sign survey, we followed trails and dry
streambeds traversing the park. The park area has three
open trail-walks: Half-hour trail, 1 h trail and 3 h trail.
These are maintained by the Forest Department. In addi-
tion, we also looked for signs on a longer 4 h trail walk
currently closed to tourists. Considering signs, we looked
for paw marks, hairs, carcasses. We followed this approach
throughout the camera-trapping days with a standard
preference to start during sunrise.
Questionnaire survey was completed with semi-struc-
tured open-ended approach. We used coloured images of
species for an ease in conversation. The aim of question-
naire survey was to get perspectives of the local Tipperah
tribe on mammalian diversity of the park area.
We used Garmin GPSMAP 64st and DNRGarmin to
obtain and handle the GPS data, ArcGIS version 10.4 (ESRI,
Redlands, USA) to process these data for geo-referenced
maps following the geo-datum WGS 1984.
Statistical analyses were carried out using R version
3.6.0 (R Development Core Team 2019). Organization of
camera trap images was done following Niedballa et al.
(2016). Plotting of activity patterns was expressed in kernel
density estimation plots. Using Ridout and Linkie (2009);
Niedballa et al. (2016); we measured activity patterns of
dhole, its potential prey species barking deer Muntiacus
muntjak and wild boar Sus scrofa in SNP. We also calcu-
lated kernel densities of anthropogenic activities on two
different categories: Human activity (appearance of human
on camera trap images) and livestock movement (appear-
ance of free-roaming cattle on camera trap images). The
Authenticated akash.muntasir/ Author's copy | Downloaded 05/14/20 03:39 AM UTC
4 T. Zakir et al.: Dhole in northeast Bangladesh
diel cycle was categorized based on sunrise and sunset as
diurnal (active during the day) and nocturnal (active dur-
ing darkness). Crepuscular time band was created by tak-
ing 1 h before and 1.5 h after sunrise and 1.5 h both before
and after sunset. Based on Gerber et al. (2012); activity
pattern was set to be cathemeral in case of irregular ac-
tivities at any time of night or day.
We estimated temporal overlap between diel activities
of dhole and those of potential prey species and anthro-
pogenic activities following Ridout and Linkie (2009).
Capture times were considered as a random sample of the
photos taken at any time of the day. The overlap data for
dhole-barking deer, dhole-wild boar, dhole-human activity
and human-livestock movement pairs were expressed us-
ing a coefficient of overlap, Δ
1 – a quantitative measure
ranges from 0 (no overlap on active periods) to 1 (complete
overlap, identical activity density). In accordance with
Ridout and Linkie (2009); we used estimator Δ
1 because it
is best to describe scenario coming with small sample sizes.
The 95% confidence intervals of these pairs were obtained
as percentile intervals from 10,000 bootstrap samples. The
MAMMALIA-2019-0050_proof  4 May 2020  9:24 pm
simulation gave the true value with actual proportion of
confidence intervals.
We followed Mackenzie et al. (2017) and ran single-
species, single-season occupancy model for dhole in SNP
using Presence version 2.12.36 (Hines 2006) to get naïve
occupancy estimate (SD/S), and detection probability (p)
for each trapping station.
We obtained a total of 32 photos of solitary individuals
from five different stations in three different days on eight
independent occasions. In comparison to the complete
effort, relative abundance index (RAI) of dhole was 1.36. To
calculate the RAI for dhole, we multiplied independent
detections by hundred, and, then divided by the total
number of camera trap nights (Azlan and Sharma 2006).
The RAI are assumed to be linearly correlated to overall
abundance in a particular area (Palmer et al. 2018). The
naïve occupancy estimate of dhole (SD/S) was 0.41 based
on detection in five out of 12 trap stations with detection
probability (p) of 0.6455.
The first picture of an individual appeared on 17
December 2018, at 0719 h (Figure 2A; Supplementary
Figure 2: Evidence of dhole Cuon alpinus in
Satchari National Park. (A) Camera Trap
Station 009 17 December 2018 at 07:19 h;
(B) Camera Trap Station 012 15 January
2019 at 09:52 h; (C) Camera Trap Station
003 15 January 2019 at 11:44 h; (D) Camera
Trap Station 008 15 January 2019 at 12:06 h;
(E) Camera Trap Station 011 17 January
2019 at 07:39 h; (F) Footprint of dhole
(photo: Harish Debbarma).
Authenticated akash.muntasir/ Author's copy | Downloaded 05/14/20 03:39 AM UTC

Table S1). We identified dholes from another four locations
later on (Figures 2B–E). It is noteworthy that, on 15 January
2019, three camera stations captured dhole.
A single individual was captured on every image.
However, we could only identify two different individuals
which appeared on station 07. On 2021 h of 19 January 2019,
within 5 s, two individuals singly appeared on camera
pacing at the same direction. We confirmed on their in-
dividuality as the camera function was set as three
continuous images and one 8 s video in response to every
detection. One individual appeared on the last two of the 3-
image series, the other on the video of the same sequence
started immediately after the photo series. For the rest of
the images, we could not identify whether it was the same
specimen or a different one.
Of all positive camera stations for dhole, the deploy-
ment ID 07 got the clearest and the highest number of
images (n = 12) on two events (Supplementary Table S1).
This was installed on a streambed that runs below a bridge
and connects the RHRF with the park. The individual
seemed curious, crossed the camera twice. We surmised
that the individual had attempted to cross the underpass
yet got startled for unknown reason and went back to the
trail where it came from. Figure 3 depicts activity of dhole
in SNP, its temporal overlap with potential prey species
and human activity. Dhole appeared to be mostly diurnal
with moderate degree of temporal overlap with barking
deer (Δ
1  0.65) human activity Δ
1  0.63) and livestock
movement Δ
1  0.59) (Figure 3A; C-D respectively).
Figure 3B indicates low overlap between dhole and wild
MAMMALIA-2019-0050_proof  4 May 2020  9:24 pm
T. Zakir et al.: Dhole in northeast Bangladesh 5
boar activity (Δ
1  0.31) as the latter being completely
nocturnal. Observing the 95% confidence interval for the
range of coverage’s for each pair, it becomes evident that
the overlap estimate for dhole and barking deer shared a
narrower range; hence, higher likelihood of sharing the
same activity period. Supplementary Table S1 narrates the
dhole encounters in detail.
During the questionnaire survey, we found claims on
the presence and sightings of dhole from the park area.
Also, the interviewees reported findings of corpses of both
barking deer and wild boar allegedly attacked by dholes.
We compiled five incidents that were evidence of dhole
activity where at least two encounters indicated livestock
depredation (Supplementary Table S1).
Our study is the first confirmed evidence of dhole
presence from northeast Bangladesh. Owing to prey
abundance and absence of any other sympatric apex
predators, we surmise that dhole likely uses the park area,
even the whole RHRF as well. This is based on direct ob-
servations as well as camera trap photos of two medium-
sized ungulate prey species: Barking deer M. muntjak and
wild boar S. scrofa. They appeared on 10 out of 12 camera
trap stations and are potential prey populations for dholes.
Kamler et al. (2012) and Grassman et al. (2005) stated
barking deer is the primary prey of dholes that live in
similar habitats in Southeast Asia. In addition to the
abundance of barking deer and wild boar, we also recorded
the presence of eight smaller carnivore and mesocarnivore
species: leopard cat Prionailurus bengalensis, fishing cat
Prionailurus viverrinus, jungle cat Felis chaus, large Indian
Figure 3: Daily activity patterns of dhole
Cuon alpinus in SNP and its temporal
overlap with potential prey species and
anthropogenic factors: (A) with barking
deer Muntiacus muntjak; (B) with wild boar
Sus scrofa; (C) with human acivity and (D)
livestock movement. The x-axis represents
the active time from all activity samples; the
y-axis range is the kernel density. The
vertical dashed (—) line pairs indicate the
crepuscular time band and separate diurnal
and nocturnal activity periods. The shaded
area under each density estimate
comparison denotes the corresponding
overlap coefficient (Δ
1 ). CI stands for the
95% confidence interval.
Authenticated akash.muntasir/ Author's copy | Downloaded 05/14/20 03:39 AM UTC
6 T. Zakir et al.: Dhole in northeast Bangladesh
civet Viverra zibetha, common palm civet Paradoxurus
hermaphroditus, masked palm civet Paguma larvata, crab-
eating mongoose Herpestes urva, golden jackal Canis
aureus and yellow-throated marten Martes flavigula indi-
cating a diverse carnivore community in the park area.
Considering barking deer as potential prey of dhole for
the region and scaling five other reserve forests of north-
east Bangladesh, we found that studies pertinent to
mammalian diversity are minimal. In this region, of all
available studies, only Feeroz et al. (2011) and Rahman
(2017) applied camera-traps in their study sites. Rahman
(2017) deployed cameras in AHRF, PHRF RRF, THRF and
WBRF whereas Feeroz et al. (2011) only to a wildlife sanc-
tuary of THRF (Figure 1). But, there is no rigorous research
in the region that deployed a prolonged and systemic
camera-trapped survey, neither there is any study on
barking deer abundance in any of the forests of Figure 1A:
Results of which may indicate viability of presence of large,
and cryptic predatory species like leopard, clouded leop-
ard, Asiatic golden cat, marbled cat, etc.
In Tripura, known dhole presence is in the Trishna
Wildlife Sanctuary and the Gumti Wildlife Sanctuary; both
are more than 100 km in the south from our study area
(Choudhury 2013). Figure 1A, interestingly, indicates that
all of the six reserve forests of Bangladesh are extensions of
the Baramura-Atharamura-Longtharai-Unakoti hill ranges
that possess forested areas. Though border fencing is an
issue to restrict dhole movement, borderline forests are still
dense in these reserve areas, particularly in the THRF and
the RRF (Figure 1A). In relevance, we discovered dhole
from an area where regular illegal logging and tourist ac-
tivities are prominent disturbances.
Taking these factors into account and as dhole is
generalist in habitat utilization (Kamler et al. 2015), we
recommend thorough surveys for these least known forests
of northeast Bangladesh. Methodical surveys may lead to
the discovery of a resident population of the species in
Bangladesh and to unique insight to their coping strategies
in degrading habitats sprawled across geopolitical
boundaries between India and Bangladesh. Such studies
are in dire need as dhole can serve as a keystone species
rejuvenating conservation for these long-neglected van-
ishing forests of northeast Bangladesh.
Acknowledgements: We express our gratitude to
Bangladesh Forest Department for granting permission
to conduct the survey in SNP. This work was funded by
the WildTeam and the Ministry of Science and Technol-
ogy, Government of the People’s Republic of Bangladesh,
both were for Tania Zakir to pursue her MSc thesis
MAMMALIA-2019-0050_proof  4 May 2020  9:24 pm
research. We received equipment assistance and tech-
nical support from WildTeam. Thanks to Mizanur Rah-
man, Fatema Tuz Zahura, Rafia Mahjabin and Zaber Khan
for companionship during the field work and to the
people of SNP who helped us on several occasions. We
are also grateful to Dr Jan Kamler, Dr Kashimra Kakati, Dr
Anwaruddin Choudhury and Dr William Duckworth for
their invaluable insights on the distribution of dhole in
northeast India.
References
Ahamed, Y.S. (1981). With the wild animals of Bengal, Dhaka. p. 84.
Azlan, J.M. and Sharma, D.S. (2006). The diversity and activity
patterns of wild felids in a secondary forest in Peninsular
Malaysia. Oryx 40: 36–41, https://doi.org/10.1017/
s0030605306000147.
Boitani, L. and Powell, R.A. (Eds.) (2012). Carnivore ecology and
conservation: A handbook of techniques. Oxford University
Press, New York, NY, p. 506, https://doi.org/10.1093/acprof:
oso/9780199558520.001.0001.
CCA. (2016). A preliminary wildlife survey in Sangu-Matamuhuri
Reserve Forest, Chittagong Hill Tracts, Bangladesh.CCA, Creative
Conservation Alliance, Dhaka, Unpublished report submitted to
Bangladesh Forest Department, p. 52.
Chakma, S. (2015). Assessment of large mammals of the Chittagong
Hill Tracts of Bangladesh with emphasis on tiger Panthera tigris.
PhD Thesis (unpubl.), Department of Zoology, University of
Dhaka, Dhaka, p. 209.
Choudhury, A. (2013). The mammals of North East India. Gibbon Books
and the Rhino Foundation for Nature in NE India, Guwahati,
p. 432.
Choudhury, J.K., Biswas, S.R., Islam, M.S., Rahman, O., and Uddin,
S.N. (2004). Biodiversity of Satchari Reserve Forest, Habiganj.
IUCN, International Union for Conservation of Nature,
Bangladesh Country Office, Dhaka, p. 130.
Durbin, L.S., Venkataraman, A., Hedges, S., and Duckworth, W.
(2004). Dhole Cuon alpinus. In: Sillero-Zubiri C., Hoffmann M.
and MacDonald D.W. (Eds.) Canids: Foxes, wolves, jackals and
dogs. status survey and conservation action plan. IUCN/SSC
Canid Specialist Group, Gland, pp. 210–219, https://doi.org/10.
1108/RR-08-2018-0126.
Feeroz, M.M., Hasan, M.K., and Khan, M.M.H. (2011). Biodiversity of
protected areas of Bangladesh, vol. 1. Rema-Kalenga Wildlife
Sanctuary. Arannayk Foundation, Dhaka, p. 219.
Gerber, B.D., Karpanty, S.M., and Randrianantenaina, J. (2012).
Activity patterns of carnivores in the rain forests of Madagascar:
implications for species coexistence. J. Mammal. 93: 667–676,
https://doi.org/10.1644/11-mamm-a-265.1.
Grassman , Jr, L.I., Tewes, M.E., Silvy, N.J., and Kreetiyutanont, K.
(2005). Spatial ecology and diet of the dhole Cuon alpinus
(Canidae, Carnivora) in north central Thailand. Mammalia 69:
11–20, https://doi.org/10.1515/mamm.2005.002.
Hines, J.E. (2006). PRESENCE 2. Software to estimate patch occupancy
and related parameters. USGS‐PWRC, Maryland. https://www.
mbr-pwrc.usgs.gov/software/presence.html.
Authenticated akash.muntasir/ Author's copy | Downloaded 05/14/20 03:39 AM UTC

IUCN Bangladesh. (2015). Red list of Bangladesh, vol. 2. Mammals.
IUCN, International Union for Conservation of Nature,
Bangladesh Country Office, Dhaka, p. 250.
Jenks, K.E., Kitamura, S., Lynam, A.J., Ngoprasert, D., Chutipong, W.,
Steinmetz, R., Sukmasuang, R., Grassman, L.I., Cutter, P.,
Tantipisanuh, N., et al. (2012). Mapping the distribution of dholes,
Cuon alpinus (Canidae, Carnivora), in Thailand. Mammalia 76:
175–184, https://doi.org/10.1515/mammalia-2011-0063.
Kamler, J.F. (2013). Extreme dogs: Dhole. Wildlife World 3: 15–16.
Kamler, J.F., Johnson, A., Vongkhamheng, C., and Bousa, A. (2012).
The diet, prey selection, and activity of dholes (Cuon alpinus) in
northern Laos. J. Mammal. 93: 627–633, https://doi.org/10.
1644/11-mamm-a-241.1.
Kamler, J.F., Songsasen, N., Jenks, K., Srivathsa, A., Sheng, L. and
Kunkel, K. (2015). Cuon alpinus. The IUCN Red List of Threatened
Species 2015: e.T5953A72477893. [online] https://dx.doi.org/
10.2305/IUCN.UK.20154.RLTS.T5953A72477893.en.
Khan, M.M.H. (2018). Photographic guide to the wildlife of
bangladesh. Arannayk Foundation, Dhaka, p. 488.
MacKenzie, D.I., Nichols, J.D., Royle, J.A., Pollock, K.H., Bailey, L., and
Hines, J.E. (2017). Occupancy estimation and modeling: inferring
patterns and dynamics of species occurrence. Elsevier,
Amsterdam, p. 648.
Mukul, S.A., Uddin, M.B. and Tito, M.R. (2006). Study on the status
and various uses of invasive alien species in and around Satchari
National Park, Sylhet, Bangladesh. Tigerpaper 33: 28–32.
Niedballa, J., Sollmann, R., Courtiol, A., and Wildting, A. (2016).
‘camtrapR’: an R package for efficient camera trap data
management. Methods Ecol. Evol. 712: 1457–1462, https://doi.
org/10.1111/2041-210x.12600.
NSP. (2006). Site status report: Satchari National Park. NSP, Nishorgo
Support Project, United States Agency for International
Development (USAID) and Bangladesh Forest Department (BFD),
International Resources Group (IRG). Dhaka, p. 4.
O’Connell, A.F., Nicholas, J.D. and Karanth, K.U. (2011). Camera traps
in animal ecology. Springer, USA, p. 263.
Palmer, M.S., Swanson, A., Kosmala, M. and Arnold, T. (2018). Packer
Evaluating relative abundance indices for terrestrial herbivores
from large‐scale camera trap surveys. Afr. J. Ecol. 56: 791–803,
https://doi.org/10.1111/aje.12566.
View publication stats MAMMALIA-2019-0050_proof  4 May 2020  9:24 pm
T. Zakir et al.: Dhole in northeast Bangladesh 7
R Development Core Team. (2019). R: A language and environment for
statistical computing. R Foundation for Statistical Computing,
Vienna, Austria. https://www.R-project.org.
Rahman, H.A. (2017). Mammal biodiversity in the northeast forests,
and the distribution of fishing cats in Bangladesh. MSc. Thesis
(unpubl.), Department of Entomology and Wildlife Ecology,
University of Delaware, Delaware, p. 100.
Ramesh, T., Sridharan, N., Sankar, K., Qureshi, Q., Selvan, K.M.,
Gokulakkannan, N., Francis, P., Narasimmarajan, K., Jhala, Y.V., and
Gopal, R. (2012). Status of large carnivores and their prey in tropical
rainforests of South-western Ghats, India. Trop. Ecol. 53: 137–148.
Ridout, M.S. and Linkie, M. (2009). Estimating overlap of daily activity
patterns from camera trap data. J. Agr. Biol. Envir. St. 14:
322–337, https://doi.org/10.1198/jabes.2009.08038.
Srivathsa, A., Kumar, N.S., and Karanth, K.U. (2017). Home range size
of the dhole estimated from camera-trap surveys. Canid. Biology.
and Conservation. 20: 1–4.
Srivathsa, A., Karanth, K.U., Kumar, N.S., and Oli, M.K. (2019). Insights
from distribution dynamics inform strategies to conserve a dhole
Cuon alpinus metapopulation in India. Sci. Rep. 9: 3081, https://
doi.org/10.1038/s41598-019-39293-0.
TEAM Network. (2011). Terrestrial vertebrate protocol implementation
manual. version. 3.1. tropical ecology, assessment and
monitoring network, Center for Applied Biodiversity Science,
Conservation International, Virginia. p. 69.
Van Valkenburgh, B. (1991). Iterative evolution of hypercarnivory in
canids (Mammalia: Carnivora): evolutionary interactions among
sympatric predators. Paleobiology 17: 340–362, https://doi.
org/10.1017/s0094837300010691.
WDPA. (2016). Satchari National Park. WDPA, World Database on
Protected Areas, United Nations Environment Programme,
Nairobi.
Wearn, O. and Kapfer, W.P. (2017). Camera-trapping for conservation:
a guide to best-practices. WWF, World Wildlife Fund, Woking,
Surrey, p. 164. https://doi.org/10.13140/RG.2.2.23409.17767.
Supplementary Material: The online version of this article offers
supplementary material (https://doi.org/10.1515/mammalia-2019-
0050).
Authenticated akash.muntasir/ Author's copy | Downloaded 05/14/20 03:39 AM UTC