2020 - Akash Et Al - Indian Gray Wolf in Bangladesh

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On the reappearance of the Indian grey wolf in Bangladesh after 70 years:


what do we know?

Article  in  Mammalian Biology - Zeitschrift fur Saugetierkunde · September 2020


DOI: 10.1007/s42991-020-00064-4

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Mammalian Biology
https://doi.org/10.1007/s42991-020-00064-4

SHORT COMMUNICATION

On the reappearance of the Indian grey wolf in Bangladesh


after 70 years: what do we know?
Muntasir Akash1 · Umar Faruq Chowdhury2 · Fatema‑Tuz‑Zohora Khaleque3 · Rifath Nehleen Reza2 ·
Dulal Chandra Howlader1 · Mohammad Riazul Islam2 · Haseena Khan2

Received: 31 May 2020 / Accepted: 29 August 2020


© Deutsche Gesellschaft für Säugetierkunde 2020

Abstract
The Indian grey wolf, Canis lupus pallipes Sykes, 1831, is a small, cryptic subspecies and the only wolf living in arid plains
and deserts of the Indian subcontinent. Since 1950, it has been considered extinct beyond 88° east longitude. Herein, we
report an instance from Bangladesh after 70 years. A solitary male of C. l. pallipes was killed in retaliation in June 2019 as
livestock predation events erupted and lasted for a month after a severe cyclone had swept coastal Bangladesh. The specimen
was about 119 cm from nose to tail tip with a skull length of 26.23 cm. Two molecular markers, mt d-loop control region
and 16S rRNA, and 54 cranial parameters consolidated the identity. Bayesian inference and maximum-likelihood analyses
indicated its intraspecies position. The locality of conflict, 450 km eastward of the easternmost population of C. l. pallipes, is
adjacent to the Sundarbans in the Ganges estuary that presents formidable tidal rivers as dispersal barriers. In 2017, another
wolf was sighted from the Indian Sundarbans vicinity. The present incident and the sighting of 2017 remarkably appeared
from the farthest corners of a 10,000 km2 strong mangrove network that is rimmed by dense human settlements. The records
surmise about the most challenging wolf dispersal route ever recorded. Additionally, the south-central coasts of Bangladesh,
once home to wolves, bear old planted mangroves with open dunes but never surveyed for mammals. These facts necessitate
a systematic camera-trapping in the coastal mangroves of Bangladesh exclusively intended for wolves.

Keywords Wolf · Canis lupus pallipes · Sundarbans · Bangladesh

Evolved in the Pleistocene period, grey wolf (Canis lupus Canidae, is one of the most successful of any contempo-
Linnaeus, 1758), the largest extant member in the family rary large carnivore mammals. The present range of wolf
is still splayed across most of Asia and North America;
also includes places in Europe and North Africa in spite of
Handling editor: Laura Iacolina. localized extinction and surmounting anthropogenic pres-
sure (Mech and Boitani 2004; Hunter and Barrett 2018).
Electronic supplementary material  The online version of this
article (https​://doi.org/10.1007/s4299​1-020-00064​-4) contains
supplementary material, which is available to authorized users.

* Muntasir Akash Haseena Khan


m17.zoo@du.ac.bd haseena@du.ac.bd
Umar Faruq Chowdhury 1
Department of Zoology, Faculty of Biological Sciences,
umarfchy@gmail.com
University of Dhaka, Dhaka, Bangladesh
Fatema‑Tuz‑Zohora Khaleque 2
Department of Biochemistry and Molecular Biology,
ftzmila@gmail.com
Faculty of Biological Sciences, University of Dhaka, Dhaka,
Rifath Nehleen Reza Bangladesh
nehleen11@gmail.com 3
Bangladesh Forest Department, Ministry of Environment
Dulal Chandra Howlader and Forests, Dhaka, Bangladesh
dulalhowlader1980@gmail.com
Mohammad Riazul Islam
mriazulislam@du.ac.bd

13
Vol.:(0123456789)
M. Akash et al.

In response to diverse habitats, wolves have become highly appeared to be the entire south-central and western Bengal
adaptive (Mech and Boitani 2004). (Fig. 1a). Hunter (1876c) also noted wolves from Darjeeling,
Wolves are accounted for as many as 36 subspecies by a northern hilly region of West Bengal (Fig. 1a); however,
Wozencraft (2005); 12 were mentioned by Hunter and Bar- whether these wolves were plainland (C. l. pallipes) or hilly
rett (2018) with skepticism. However, with the advent in subspecies (C. l. chanco) was not mentioned. Mitra (1957)
molecular phylogenetics, different lineages of wolf have noted a predation event of wolf killing a man around 1940s
been recognized; many of which are elevated to species from the coastal grasslands of Noakhali, south-central Bang-
level as for the African wolf (Canis lupaster Hemprich and ladesh (Fig. 1a). O’Malley (1914,1923) mentioned wolf
Ehrenberg, 1832) and wolves of southeastern North America from the districts of Noakhali and Pabna of Bangladesh as
(eastern wolf C. lupus lycaon Schreber, 1775 and red wolf well as from Murshidabad, Malda, and Dinajpur—districts
C. l. rufus Audubon and Bachman, 1851) (Vonholdt et al. which are now part of India (Hunter 1876a,b) (Fig. 1a).
2016; Sinding et al. 2018; Alvares et al. 2019). Origin of the These regions lie along the Ganges River and rich in river-
divergence in the Holarctic grey wolves (Canis lupus spp.) ine open country and grassland mosaics (IUCN Bangladesh
is tracked down to South Asia (Shrotriya et al. 2012). The 2015) similar to current range areas of the Indian wolves.
only two South Asian subspecies of grey wolves, the Hima- However, long before the independence of Bangladesh,
layan wolf (Canis lupus chanco Gray, 1863) and the Indian wolf is regarded as an extirpated species in the eastern Ben-
wolf (Canis lupus pallipes Sykes, 1831), also the most basal gal (Khan 2015, 2018), although any exclusive survey has
ones, are accounted for the global spread of wolves (Shro- never been undertaken.
triya et al. 2012; Sharma et al. 2019). Interestingly, these two This work proved the appearance of the Indian grey wolf
lineages were suspected as distinct species by Sharma et al. in Bangladesh after 70 years. Craniometric measurements
(2004) and Aggarwal et al. (2007). Werhahn et al. (2020) are also provided for the first time from the entire Indian
stressed further taxonomic recognition for the more ancient Subcontinent. Its phylogenetic position is also examined.
Himalayan wolf. Finally, the possible reasons causing its appearance in the
Compared to the range-restricted C.l. chanco, C. l. pal- country are discussed.
lipes is small and occupies a wider distribution (Prater On 4th June 2019, an animal allegedly deemed a wolf,
1971; Menon 2014). This subspecies is almost half of the was killed in retaliation at a coastal village of Bangladesh
size when compared to the Holarctic congeners (Hunter (N 22.01374 E 90.1566). The locality, administratively, falls
and Barrett 2018). Roughly equal to an oversized golden under the district of Barguna (Fig. 1b). The area is jutting
jackal (Canis aureus Linnaeus, 1758), coat pattern makes into the Bay of Bengal, traversed by three coastal rivers of
the Indian wolf cryptic and less evident (Prater 1971; the Ganges drainage system. The southern rim of the dis-
Menon 2014; Hunter and Barrett 2018). Canis lupus pal- trict is bordered by mangrove belts (Fig. 1b). A visit was
lipes is known as a habitat-generalist occupying dry and made to the place on 12th June 2019. Direct morphological
semi-arid habitats such as plains, grassland-sandbar, and measurements could not be taken as the animal was buried
scrubland–agricultural mosaics (Jhala 2003; Menon 2014; on the same day of the killing on the premises of the local
Saren et al. 2019). There is no range-wide assessment of the outpost (N 21.9848 E 21.9848) of the Bangladesh Forest
Indian wolves as in for the European and the Mediterranean Department. However, for molecular analyses, muscular tis-
wolves (Jdeidi et al. 2010; Boitani 2018). In India, about sues from the upper left arm of the specimen were collected
2000–3000 individuals are believed to be present; however, after exhumation and instantly preserved on 95% ethanol.
studies on the Indian wolves appeared in a patchy pattern Through semi-structured interviews, local news correspond-
(Jhala 2003; Sharma et al. 2019). ents, forest personnel, and villagers were asked about the
The extant range for the Indian wolf, though patchy and general appearance of the animal and its conflict records.
fragmented, goes from the Peninsular India to the Arabian For the geographic information on the locality, Denzau et al.
deserts (Jhala 2003). However, it is not known beyond 88° (2015), IUCN Bangladesh (2015), and Islam et al. (2020)
east longitude, except for a recent sighting from a Sundar- were followed as references. The pertinent maps were ren-
bans outskirt in the state of West Bengal, India (Mukherjee dered using WGS 1984 geo-datum and ArcGIS 10.5.
2017) (Fig. 1a). Northerly, the subspecies is now contained On 13th December 2019, the skeletal remains were again
by the Ganges River in the state of Bihar, India (Dey et al. exhumed. Hafner et al. (1984) and Rowley (2015) were used
2010; Sharma et al. 2019) (Fig. 1a). Anecdotes on C. l. pal- as references for skull preparation. After degreasing and
lipes from the historic region of Bengal, now succeeded cleaning with detergents and 3% commercial-grade solution
by the state of West Bengal, India, and Bangladesh, went of hydrogen peroxide, the sun-dried skull was polished with
back to the 1950s (Fig. 1a). Through accounts of Hunter a soft brush. Cyanoacrylate adhesives and silicone sealants
(1876a,b,c), Simson (1886), and O’Malley (1914, 1923), the were applied in case of shattered skull fragments. With a
historic eastern limit of wolves in the Indian subcontinent 30 cm-long slide calipers, measurements were noted to an

13
On the reappearance of the Indian grey wolf in Bangladesh after 70 years: what do we know?

Fig. 1  Occurrence of the Indian


grey wolf Canis lupus pallipes
in its eastern limit (a) and the
Sundarbans forest peripheries
(b). Black dashed line marks
historical easternmost limit of
the species after Simson (1886).
Green-shaded areas denote
forest cover. Light grey areas
denote the historic region of
Bengal, now succeeded by the
state of West Bengal, India, and
Bangladesh. Dark grey area
denotes the district of Barguna.
Inset shows the region within
the Indian subcontinent

accuracy of 0.01 mm. For craniometric parameters, 54 dif- the standard procedure of the phenol–chloroform protocol
ferent measurements were noted (Supplementary Material was followed and incubation at − 20 °C was carried out
Fig. B1) following Clutton-Brock et al. (1994) and Krizan overnight. The isopropyl alcohol precipitation method was
(2005). All measurements were taken four times each (two then conducted; DNA precipitate was re-suspended in 60 μl
by MA and two by DCH). Then, the mean value for each nuclease-free water. Amplification of two molecular mark-
parameter was used. Dental anomalies and abrasions were ers was targeted: 16S rRNA (16S) and mitochondrial (mt)
observed according to Janssens et al. (2016). D-loop control region. To design the primers for polymerase
To extract DNA, 20 mg sample was mixed into 100 μl of chain reaction (PCR), sequences for 16S and mt D-loop con-
TESU6 buffer and incubated at 55 °C for 15 min in shaking trol region of grey wolf (Canis lupus) were retrieved from
incubator (Aranishi and Okimoto 2006). Ten μl of 5.0 M the National Center for Biotechnology Information (NCBI)
sodium chloride (NaCl) was added to the solution. Then, GenBank database and were aligned using Clustal Omega

13
M. Akash et al.

(Madeira et al. 2019) to find the highly conserved regions. Bayesian Information Criterion (BIC) values. Priori settings
Parameters of the candidate primers were checked by IDT were applied with BEAUti v. 1.8.0 (Drummond et al. 2012).
Oligo Analyzer (Owczarzy et al. 2008) and Primer Blast A Tamura-Nei (TrN) model using invariant sites and gamma
(Ye et al. 2012). Finally, for mt d-loop control region, the (I + G) was applied (Tamura and Nei 1993) with five gamma
following primers were used: 5′-TGA ATC ACC CCT ACT categories and the assumption of a strict clock. All default
GTG-3′ (forward) and 5′-CCA TTG ACT GAA TAG CAC settings were maintained and a random tree was selected for
C-3′ (reverse). For 16S primers, 5′-GGA GCG ATA GAG starting; only the Tree Prior category was set to the Yule Pro-
ATA GTA CC- 3′ (forward) and 5′-GTG GTC TAT CCG cess (Gernhard 2008), as for species-level analyses. BEAST
TTC CTG-3′ (reverse) were used. According to Palumbi v. 1.10.4 (Drummond et al. 2012) was used to conduct three
et al. (1991), PCR in 10 μl reaction volume was carried out independent runs of 1­ 07 generations with the dataset consid-
containing 1X standard Taq reaction buffer, 2.5 mM mag- ering the first 10% as burn-in. TRACER v. 1.7.1 (Rambaut
nesium chloride, 60 pMol primer, 3 pMol deoxynucleoside et al. 2018) was used to assess the accuracy of the output.
triphosphate (dNTP), one unit Taq DNA polymerase, and LogCombiner v. 1.10.4 was used to merge the results; the
58–59 ng DNA. The amplification was done under the fol- consensus tree was then constructed using TreeAnnotator v.
lowing conditions: 5 min of initial preheating at 95 °C; 35 1.10.4 to observe the Bayesian posterior probabilities (BPPs)
cycles of denaturation at 95 °C for 30 s; 40 s annealing at as suggested by Drummond et al. (2018). Figtree v. 1.4.4
55 °C for mt d-loop, and 57 °C for 16S; extension for 40 s (Rambaut and Drummond 2018) was used to visualize and
at 72 °C and a final extension at 72 °C for 7 min followed edit the consensus tree. Also, maximum-likelihood (ML)
by cooling at 4 °C. The PCR product was run on 1% aga- phylogenetic tree was constructed using MEGA X (Kumar
rose gel electrophoresis and visualized in Geldoc (E-Box, et al. 2018) with 1000 bootstrapping replications. The final
Vilber Smart Imaging) (Supplementary Material Fig. A1). topology of the tree was based on superior log-likelihood
Sequencing was done at the National Institute of Biotech- value.
nology, Bangladesh. The forward and reverse sequences of Our work on its genetics, craniometric measurements, and
each PCR product were assembled into contigs using CAP3 photo evidence specified the animal as a male Indian wolf
server (Huang and Madan 1999). The sequences have been (C. l. pallipes) (Figs. 2, 3; Supplementary Material Fig. B2).
submitted at NCBI GenBank database, available with acces- Uniformly thin pelage with less underfur, long muzzle and
sion number: MN219436 and MN219437. BLASTn search relatively longer legs gave the animal a lighter built. The
was done against the standard non-redundant nucleotide base color was grey mixed with black; however, flanks and
database for both the sequences to ascertain the taxonomic undersides were buff to nearly white. It was reportedly about
status of the species (Morgulis et al. 2008). 119 cm from nose to tail tip, 91.5 cm in head–body length,
To understand inter- and intraspecies genetic distance of and 71.12 cm at shoulder height (Prater 1971; Menon 2014)
the specimen, mt D-loop control region gene fragment was (Supplementary Material Fig. B2). The skull was 26.23 cm
considered and 17 relevant sequences were retrieved from in total length and 23.62 cm in condylobasal length. Sup-
NCBI GenBank database. Sequences submitted from the plementary Material B (Table B.1) presented all 54 skull
Indian wolf range area were considered, i.e., four of Sharma parameters, whereas definitions of these traits were por-
et al. (2004) and five of Aggarwal et al. (2007) entitled as trayed in Fig. A.1. The measurements were indicative of
C. l. pallipes and of C. indica, respectively. Five sequences a male specimen of the Indian wolf according to Menon
of the Himalayan wolf were included: Accession no. (2014) and Khosravi et al. (2014).
KY94030.1, KY996529.1, and KY996530.1 were named as Cranial damages and anomalies were conspicuous. Skull
Canis himalayensis by Werhahn et al. (2017); KT321360.1 trauma owing to mob beating was evident (Fig. 2a). The
and AB007379.1 as C. l. chanco by Chetri et al. (2016) right cheekbone was shattered; parts from of zygomatic
and Tsuda et  al. (1997) accordingly. The Eurasian wolf arch along the squamosal suture were missing. Traumatic
(C. l. lupus) (KY550013.1) and stray dog C. l. familiaris abrasion was observed on the right orbital process of the
(MF185437) were also considered in the analysis, sourced frontal bone. Fractures were present on nasal and maxilla.
from Montana et  al. (2017) and Strakova et  al. (2016), On the third premolar (p3) of left maxilla, an enamel chip
respectively. Golden jackal Canis aureus (MF185456) was was missing; the fracture was very likely to blunt trauma
used as outgroup (Yumnam et al. 2015) (Supplementary (Fig. 2f). Congenital dental abnormality was noted follow-
Material Table A.1). ing Janssens et al. (2016). Agenesis was observed: the first
All the sequences were aligned in MEGA X (Kumar premolar (p1) on left mandible was absent without any trace
et al. 2018) using the MUSCLE (Edgar 2004) alignment of alveolar. Medial displacement was also present on left
algorithm. ModelGenerator v. 0.85 (Keane et al. 2006) was mandible: the following premolar (p2) and its alveolar were
used to examine the best fitting model of substitution by deformed (Fig. 2g). The dentition, except the hypodontia on
observing the Akaike Information Criterion (AIC) and the left mandible, was of the normal pattern for wolf i.e., incisor

13
On the reappearance of the Indian grey wolf in Bangladesh after 70 years: what do we know?

Fig. 2  Phylogram of the Indian


grey wolf Canis lupus pallipes
and other closely related wolf
subspecies inferred from Bayes-
ian analysis. Placement of the
wolf specimen from Bangladesh
is highlighted in grey. Numeric
above the nodes denotes Bayes-
ian posterior probabilities (BPP)
on left and maximum-likelihood
(ML) bootstrap values on right.
Dashes (−) remark BPP < 0.75
or ML bootstrap support < 60.
Detail information of sequences
is provided in supplementary
material table A.1

(3/3), canine (1/1), premolar (4/4), and molar (2/3) (Görner it ends up forming the largest single block of the mangrove
and Hackethal 1987). forest, the Sundarbans. At least, 15 large estuarine rivers
Blast analysis of the sequences obtained from both PCR traversed the entire 10,000 km2 forest dividing into islands
products supports the identity as wolf Canis lupus (Sup- of varying sizes (Denzau et al. 2015; Khan 2018). Intercon-
plementary Material Fig. A. 2). Based on the phylogenetic nected by canals, creeks, and rivulets, the tidal rivers run
analyses using both maximum-likelihood and Bayesian nearly vertical in a north–south direction lying eastward in
inference methods, the wolf of Bangladesh belongs to a parallel from West Bengal up to the locality of occurrence
unique monophyletic clade together with the Indian wolf (Denzau et al. 2015) (Fig. 1a; Supplementary Material C).
sequences (Fig. 2). The clade included all the submissions On a second note, the adjacent districts of the Sundarbans
of both Sharma et al. (2004) and Aggarwal et al. (2007) periphery hold dense anthropogenic settlements both in the
together supported by BPP/ML bootstrap value of 1.00/82. state of West Bengal, India, and Bangladesh (Supplemen-
Though NCBI GenBank entries were entitled as C. indica, tary Material Table C.1). Except mangrove, these districts
the Indian grey wolf is globally accepted as C. l. pallipes have no other forest cover. The anthropogenic landscape
(Wozencraft 2005); hence, both C. indica and C. l. pallipes is dominated by fisheries farms (IUCN Bangladesh 2015)
entries of NCBI GenBank refer to the wolf subspecies of the rather than agroforestry, a known and suitable habitat for
Indian plains. The individual of Bangladesh did not show wolves (Jhala 2003; Menon 2014). For wolves, these riv-
any significant phylogenetic distance (supported by BPP/ ers appear as potential barriers. Because, the effects of the
ML bootstrap values) with the sequences of Sharma et al. Sundarbans rivers on tigers (Panthera tigris), despite living
(2004) and Aggarwal et al. (2007). in the Sundarbans for centuries, are also proven significant
The locality in Bangladesh and the known distribution of (Aziz et al. 2018).
the Indian wolf seemed remarkably instigating. As of Saren Additionally, we found that taxonomy and ecology of the
et al. (2019), West Midnapore district of West Bengal, India cryptic Indian grey wolf, throughout its known limit, are
holds the easternmost population of wolves and, at a linear still least understood. Of particular importance to note is
distance, about 450 km away from Barguna (Fig. 1a). In that, in 2017, another sighting of a male wolf at an edge
contrast to the long-ranging behavior in wolf described by of the Indian Sundarbans also created a stir similar to the
Mech and Boitani (2004), Scurrah (2012), and Sharma et al wolf of Bangladesh (Mukherjee 2017). The incident was
(2019), the region has an array of barriers which are absent concluded as a dispersal behavior though the sighting spot, a
in other wolf range countries and challenges dispersal move- village grove, is severed from the Indian Sundarbans only by
ment of any large carnivore, let alone wolf (Fig. 1a). First, a canal and the known population is about 300 km away hin-
belonging to the Lower Ganges Delta, the region is fed by dered by the estuary of the Hooghly River and dense human
a vast network of the Ganges distributaries (IUCN Bangla- settlements (Fig. 1a). Sightings of the Indian wolf within
desh 2015; Khan 2018). At the southernmost extremities, 2 years from two of the farthest corners of the Sundarbans

13
M. Akash et al.

Fig. 3  Skull of the Indian


grey wolf Canis lupus pal-
lipes reported in this article.
Cranium: dorsal view (a),
frontal view (b), palatal view
(c), nuchal view (d), lateral
view from right (e), and lateral
view from left (f). Mandible:
dorsal view (g), frontal view
(h), lateral view from right (i)
and lateral view from left (j).
Ak akrokranion, An angular
process, Au auditory bulla, B
basion, C canine, Co condyle,
Cr coronoid process, Ent enter-
orbitale (interorbital), F frontal,
N nasion, I incisor, J jugal, M
molar, Mx Maxilla, O occipital
condyle, P premolar, Pr pros-
thion, Pt parietal, Rh rhinion,
S synsphenion, Sm squamosal,
Sc sagittal crest, St staphylion,
Z zygomatic arch of squamosal.
Numeric placed against the
corresponding tooth denotes
their corresponding number in
the order of appearance. Red-
dashed circles denote anomalies
and blunt traumas

only manifests further inquiry to understand possibly the geopolitical boundary between India and Bangladesh, and
most challenging dispersal route for wolves. Compared to the Rupsa River of Bangladesh that harbors the seaport of
the observations on wolf movement by Joly et al. (2019) and Mongla (Supplementary Material Fig. C1).
Mech and Boitani (2010), the distance might seem incon- Our surmise gets buttressed by the pattern of conflict
spicuous, however, studded with large rivers; the peripher- incidents that occurred before the killing. Attempts and
ies comprise dense human habitations. Ciucci et al. (2009) predations on poultry and livestock received national atten-
denoted a somewhat similar remark on wolf movement from tion (Supplementary Material Table C.2) only after the
the Apennines to the Alps traversing highways and settle- Extremely Severe Cyclonic Storm (ESCS) Fani swept the
ments. On the contrary, the topography of the Sundarbans area on 3rd May 2019, a month before the killing took place
and adjacent regions is very different than the documented (Joint Typhoon Warning Center 2019). The first attack of
dispersal routes of wolves. Thus, we oppose the likelihood the wolf was reported on 16th May 2019, featured by three
of any wolf individual to cross the entire rim of the Sundar- different news portals (Supplementary Material Table C.2).
bans through the peripheral human settlements. It is highly After a week, the attack on livestock became a national con-
unlikely without catching any attention; impossible at the cern, broadcasted in multiple media formats (Supplementary

13
On the reappearance of the Indian grey wolf in Bangladesh after 70 years: what do we know?

Material Table C.2). All reports showed a calf, completely est Department. The skeletal remains, after examination, stay under
half-eaten from the rear—a stark sign of a canid attack the custody of the Amtali Range Office, an outpost of the Bangladesh
Forest Department.
although initially subjected to tigers. When approached,
every respondent (n = 12) agreed on the trends of conflict.
The timeline of the cyclone and the initiation of conflict
records suggested a possible displacement of the individual
from the neighboring mangroves. References
Topographically, the mangrove systems of the Bangladesh
Sundarbans hold 32.4% grassland mosaics (2061.22 km2) as Aggarwal RK, Kivisild T, Ramadevi J, Singh L (2007) Mitochon-
drial DNA coding region sequences support the phylogenetic
of 2006 (Islam 2014). These open grass meadows support distinction of two Indian wolf species. J Zool Syst Evol Res
rich prey base composed of spotted deer (Axis axis Erxleben, 45(2):163–172
1777) and wild boar (Sus scrofa Linnaeus, 1758) (Khan and Akash M, Zakir T (2020) Other than tiger: Appraising mammalian
Chivers 2007). The entire coasts of Bangladesh bear planted carnivore studies in Bangladesh. J Threat Taxa submitted
Alvares F, Bogdanowicz W, Campbell LAD, Hatlauf J, Godinho, R,
mangrove growths with open dune system; many of which Jhala YV, Werhahn G (2019) Old World Canis spp. With taxo-
are known for wild boar and spotted deer. For instance, man- nomic ambiguity: Workshop conclusions and recommendations.
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Aranishi F, Okimoto T (2006) A simple and reliable method for DNA
bans in dimension (263.04 km2). These mangroves possess extraction from bivalve mantle. J Appl Genet 47(3):251–254
agro-grassland landscape on buffer zone (Islam et al. 2020), Aziz MA, Smith O, Barlow A, Tollington S, Islam MA, Groombridge
though large carnivores and conflict instances are unheard of JJ (2018) Do rivers influence fine-scale population genetic struc-
since the 2000s (Denzau and Denzau 2010). However, the ture of tigers in the Sundarbans? Conserv Genet 19(5):1137–1151
Boitani L (2018)  Canis lupus  (errata version published in
coastal mangroves of Bangladesh have never been exclu- 2019).  The IUCN Red List of Threatened Species 2018:
sively surveyed for small and mesocarnivores. The Sunda- e.T3746A144226239. https​://doi.org/10.2305/IUCN.UK.20182​
rbans is systematically surveyed for tigers only (Akash and .RLTS.T3746​A1196​23865​.en. Accessed 22 April 2020
Zakir 2020). Boitani L, Phillips M, Jhala Y (2018) Canis lupus. The IUCN Red
List of Threatened Species 2018: e.T3746A119623865. https​://
In conclusion, we do not overthrow the possibility of doi.org/10.2305/IUCN.UK.2018-2.RLTS.T3746​A1196​23865​
long-ranging dispersal behavior of wolves. Given the geo- .en. Accessed 22 April 2020
graphical features of the region, cryptic nature of the spe- Ciucci P, Reggioni W, Maiorano L, Boitani L (2009) Long-distance
cies, and pattern of the occurring incidents, we, through dispersal of a rescued wolf from the northern Apennines to the
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