sign in sign up
<<
Home , Broadfin shark, Ganges, Ganges Delta, Ganges shark, River shark, Sundarbans

Vol. 40: 65–73, 2019 ENDANGERED RESEARCH Published October 17 https://doi.org/10.3354/esr00981 Endang Species Res

OPENPEN ACCESSCCESS

New records of the Critically Endangered Glyphis gangeticus in Bangladeshi waters: urgent monitoring needed

Alifa Bintha Haque1,2,*, Sudipta Arka Das3

1Department of Zoology, University of , Dhaka 1000, 2Department of Zoology, University of Oxford, 11a Mansfield Rd, Oxford OX1 3SZ, UK 3Department of Biochemistry and Molecular Biology, University of Dhaka, Dhaka 1000, Bangladesh

ABSTRACT: The different species of elasmobranch found in Bangladeshi fisheries and markets were quantified in a survey between February 2016 and November 2017. This resulted in records of the Critically Endangered Glyphis gangeticus (Müller & Henle, 1839) in the waters of Bangladesh. Three records from the landing sites and shark processing centres of Cox’s Bazar in southeast Bangladesh were identified and confirmed as G. gangeticus by sequencing the expression of the cytochrome c oxidase subunit I (COI) gene and by the morphological identifica- tion of 1 individual. These represent the most recent records of this species in Bangladesh in over a decade, and only the second set of records from anywhere within the range of this elusive shark. Capture dates and traders’ knowledge of these catches suggest a year-round distribution, - ting a population within the Bay of and adjacent coastal . Although the species is pro- tected under Bangladeshi law, enforcement of the law is inadequate. Monitoring for this, and other, rare and threatened species at landing sites, rigorous surveys to locate any existing popula- tions, trade regulation, and enforcement of conservation legislation should be considered priori- ties for effective conservation.

KEY WORDS: · Marine conservation · DNA barcoding · Distribution · Elasmo- branch · Elusive shark

1. INTRODUCTION , one of the most enigmatic groups of sharks in the world (Li et al. 2015, White et al. 2015). However, The Bay of Bengal harbors a rich community of there is a paucity of information on their specific dis- marine (Amaral et al. 2017). However, little tribution in the coastal areas and tropical rivers. is known about the elasmobranchs (sharks and rays) Three species of have been reported in in this (Hoq et al. 2011) owing to a lack of spe- Bangladesh, 2 of which have been described (Glyphis cies-specific research regarding their ecology, biol- glyphis and G. gangeticus; Roberts 2006, Hoq et al. ogy, , trade and species composition (Fischer 2011), and one of which is still undescribed and et al. 2012). Moreover, the world’s largest halophytic genetically distinct (Li et al. 2015, Amaral et al. 2017). , the Reserve Forest This undescribed species, overlapping in its distribu- (SRF), is located in the west of the Bay of Bengal and tion with G. gangeticus (Amaral et al. 2017), is only is a cradle for many unique species (Gopal & Chau- known to be found in the Gangetic delta (Pal et al. han 2006), including threatened and evolutionary 2014). Within the western Indo-Pacific, the Ganges distinct species. This is a suitable habitat for river shark (G. gangeticus) has a widespread but patchy

© The authors 2019. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 66 Endang Species Res 40: 65–73, 2019

distribution (Jabado et al. 2018) (Table 1). Identifica- reported in 2006 (Roberts 2006) from Bangladesh. tion is based primarily on the jaws because whole Inconsistent monitoring resulted in long intervals individuals have rarely been found and examined. between encounters. Hence, monitoring such rare Previously believed to be restricted to freshwater in and threatened species requires multiple methods, the Ganges, this species is predominantly euryhaline, including market and landing observations, evaluat- with marine records documented by Roberts (2006). ing ecological knowledge of fishers and traders, G. gangeticus is characterized by conflicting in - habitat surveys, and the use of molecular techniques formation spanning from the early 1800s to recent such as DNA barcoding. studies. Information varies from it being designated Two pieces of regional legislation protect endan- as a species endemic to the freshwater of the Ganges gered species in the Bay of Bengal and apply to G. to it being a man-eater (Roberts 2006). While it was gangeticus. Since 2001, G. gangeticus has been mentioned to have been the ‘most common’ ground protected under Schedule I, Part II A of the Indian shark species in the Bay of Bengal (Roberts 2006), it (Protection) Act, 1972, and also under is also designated as Critically Endangered by the Schedule I of the Wildlife (Conservation and Secu- IUCN Red List of Threatened Species (Compagno rity) Act, 2012, in Bangladesh. The effectiveness of 2007), mainly because of the continued decline of these regulative measures is limited, for the fol- mature individuals, its restricted range, , lowing reasons. Firstly, effective elasmobranch increased human use of rivers, pollution and habitat landing monitoring programs run by government degradation (Compagno 1997). Moreover, it is only agencies are either absent altogether or highly known historically from museum specimens (Com- faulty. This leads to limited knowledge of threat- pagno 2007), where cryptic similarities to other spe- ened and data deficient species. Secondly, there is cies and lack of specimen data pose great challenges a lack of awareness among fisherfolk and traders to accurate morphological identification. Prior to the about the law and what they should do with pro- taxonomic resolution by Li et al. (2015), G. siamensis tected species. The present study thus aimed to and G. fowlerae were considered to be synonymous monitor the largest landing site for marine fish, to G. gangeticus, further complicating the identifica- Cox’s Bazar, Bangladesh, for 3 mo following the tion process. discovery of G. gangeticus among the elasmo- After it was first described in 1839, the species branch products traded there. was not recorded between 1867 and 1996, with the The present study provides an initial understand- 1996 record being unconfirmed (Compagno 2007). ing of the landing patterns of rare and threatened However, several specimens of Ganges sharks were elasmobranch species in Bangladesh and identifies

Table 1. Summary of historical and recent records of Glyphis gangeticus showing patchy encounter rates after long intervals. TL: total length; –: not known

Specimen No. of Location of Time of Reference Notes specimens collection collection

Historical samples 3 Lower reaches of the 19th Compagno (1997), Preserved in museums of G. gangeticus Ganges-Hooghly century Roberts (2006) in Paris, Berlin, and freshwater river system G. gangeticus − Ganges River 1996 Compagno (2007) After searching for 10 yr Further confirmation Upstream of Hooghly − Compagno et al. By a photograph of the jaw of G. gangeticus − River, at Mahishadal (2008) More recent Several Lower Sundarbans, 2006 Roberts (2006) Identified as a common encounter south of , shark, contrary to of G. gangeticus Bangladesh existing beliefs First record of 1 Mumbai, India Feb Jabado et al. (2018) Female, 266 cm TL G. gangeticus in 2016 the First record of 3 Cox’s Bazar, Feb−Nov Present study Male, 118.11 cm TL G. gangeticus in Bangladesh, 2017 (1 of the 3 specimens) Bangladesh after Bay of Bengal more than a decade Haque & Das: Ganges shark in Bangladesh 67

challenges for their conservation. It includes the 2.3. Informal interviews very first records of G. gangeticus from Bangladeshi waters since 2006. It also tests the usefulness of To quantify local knowledge of native carcharhinid incorporating local knowledge on trade, seasonality, sharks, the fishers and traders were asked to partici- and perceptions of sharks and shark conservation pate in a brief informal interview following the ex - legislation. traction of DNA samples. The interviewees were shown photographs of several species from the order (e.g. Ganges shark, pig eye shark, 2. MATERIALS AND METHODS , shark, , spot-tail shark, scalloped hammerhead and ) to de - 2.1. Sampling site, genetic sampling and analysis termine whether locals are able to differentiate them.

Cox’s Bazar was chosen as a sampling site as it is the largest landing site for marine fish by volume 3. RESULTS and is a trade hub for elasmobranchs in southeast Bangladesh. The shark landing site (n = 1) and pro- 3.1. Morphological identification cessing centres (n = 6) in Cox’s Bazar were visited twice a month (Fig. 1) between February 2016 and On 13 November 2017, a whole specimen of Gly - November 2017. Morphological data on all landed phis gangeticus (S1) was identified from the Bang la - whole elasmobranch species and 20−30 tissue sam- desh Fisheries Development Corporation (BFDC) ples of meat, fin, and muscle from jaws and skin landing site, Cox’s Bazar (Fig. 1). The identification of were collected randomly from the landing site. the specimen (Fig. 2A,B) was based on the combina- Where it was possible to differentiate individuals tion of the characteristics described in Compagno and individual species, 3 to 10 genetic samples (2001), Ebert et al. (2013), and Jabado et al. (2018). were also collected from different species or body The specimen was a male and had relatively small parts; this was to avoid redundancy, i.e. collecting eyes; a short, broad and rounded snout; no interdor- tissue from the same specimen. They were pho- sal ridge; the 1st originated from the rear tographed at the processing centres during each ends of the pectoral bases; the 2nd dorsal fin was field visit. Collected samples were preserved in almost half the height of the 1st dorsal fin; and the 98% ethanol and transported to the laboratory, anal fin had a deeply notched posterior margin where they were stored in a freezer at −20°C until (Table S1 in the Supplement). Additional information analysis. DNA barcoding with cytochrome c oxidase on teeth could not be collected as the specimen was subunit I (COI) was used to confirm the identifica- removed rapidly by the traders and buyers at the tion of Glyphis gangeticus using the methods of landing site. The specimen was probably a sub-adult Ward & Holmes (2007), Ward et al. (2008), Moura given its size (TL = 118.11 cm; measured over a et al. (2008) and Naylor et al. (2012). Samples were straight line along the axis of the body from the tip of compared to a genetic database (GenBank) using the snout to the posterior tip of the upper lobe of the blasting to match these samples with previously caudal fin in its natural condition), and the claspers collected, documented and submitted G. gangeticus were not totally calcified as determined by holding samples. A family tree with other specimens was the specimen tightly. A summary of the whole indi- then constructed. vidual and the other 2 specimens are given in Table 2. During this study, 13 individuals were photographed that were morphologically similar to Ganges sharks 2.2. Morphological measurements (Glyphis spp.). These species were investigated for a better understanding of their occurrence and to mini- Morphometric measurements and photographs of 3 mize the risk of misidentification. However, 2 were Ganges sharks were collected using the methodol- proven to be bull sharks and, due to lack of quality ogy of Compagno (2001) for all specimens (S1, S2, photographs and genetic material available, the re- and S3). The total length (TL) and mass of 2 speci- maining individuals could not be identified confi- mens (S2 and S3) were estimated from the descrip- dently. This demonstrates the challenges of identify- tion of the trader using an informal interview tech- ing Ganges sharks only from photographs while the nique (see the Supplement at www.int-res. com/ researcher is not in the field and when good quality articles/suppl/n040p065_supp.pdf ). reference photographs are not available. 68 Endang Species Res 40: 65–73, 2019

Fig. 1. Area of occurrence of Glyphis ganteticus (inset). Yellow lines indicate the range (extant) according to the IUCN, Jabado et al. (2018), Roberts (2006) and Compagno (2002). Black star: one of the sites (Cox’s Bazar) where regular landing monitoring was conducted

3.2. Identification through DNA barcoding G. fowlerae are considered to be synonyms of G. gangeticus (Li et al. 2015). The 3 specimens of G. The partial COI gene sequence of specimen 1 gangeticus cluster within the clade of G. gangeticus (GenBank accession number MH841976) was 656 bp sequences derived from individuals sampled from in length. It was blasted against sequences in GenBank various locations on the subcontinent. and there was 100% pairwise identity of the G. gangeticus sequences to GenBank accession number KT698058 (G. gangeticus isolate GN1188 mitochon- 3.3. Insights into catch location, drion, complete genome; PMID: 26460025). The par- trade, and seasonality tial COI gene sequences of specimens 2 (GenBank accession number MH841975) and 3 (MH841977) All 3 specimens were non-discarded bycatch from were also 656 bp in length. There was 99% pairwise the Bay of Bengal, Bangladesh region (not more than identity of the G. gangeticus se quences to accession 40 m depth) from artisanal fishers using an array of number KT698054 (G. gangeticus isolate GN2669 drift gillnets, set-bag nets, trammel nets, and long mitochondrion, complete genome; PMID: 26460025) lines. The fishers target a variety of marine fish, pre- when blasted against sequences in GenBank. dominantly Ilisha (Hilsa taneolosa, family Clupei- The resultant phylogenetic tree from the neigh- dae). Artisanal originate from Cox’s Bazar and bour-joining analysis of the COI data derived from go on to catch fish on a 7- to 12-d trip. These artisanal the 3 G. gangeticus specimens, together with previ- boats catch fish throughout the coastal waters ously available sequences of other representatives of between 0 and 40 m depth (Shamsuzzaman et al. G. gan geticus, G. siamensis, G. fowlerae, G. garricki 2017) and sometimes beyond. Two of the samples and G. glyphis, is shown in Fig. 3. G. siamensis and were identified from randomly selected dried and Haque & Das: Ganges shark in Bangladesh 69

Fig. 2. (A,B) Dorsal and ventral view of specimen 1 (S1) of Glyphis gangeticus, estimated total length 118 cm. (C) Semi-dried pectoral fins of specimen 2 (S2) of G. gangeticus and (D) semi-dried fins of G. gangeticus specimen 3 (S3). Estimated total length 188 cm: (a) first dorsal fin; (b) lower caudal-fin lobe; (c) left pectoral fin; (d) right pectoral fin; caught from the territorial waters of Bangladesh and landed in Cox’s Bazar semi-dried fin samples in the shark processing cen- tional trade in these species. The specimens were tres, where the majority of the dried meat is exported collected in the months of February (early summer), to (Haque et al. 2018). Therefore, these April (summer), and November (winter), indicating a samples provide evidence of the existing interna- nearly year-round catch. 70 Endang Species Res 40: 65–73, 2019

Table 2. Summary of the 3 Glyphis gangeticus specimens recorded from Cox’s Bazar, Bangladesh. TL: total length

Specimen Local name Location Date of observation TL (cm) Sex Part

S1 Bhota, Boli hangor BFDC landing site, 13 Nov 2017 118.11 Male Whole Cox’s Bazar S2 Unknown Shark processing centre, 29 Apr 2017 Unknown Unknown Pectoral fin Cox’s Bazar S3 Unknown Shark processing centre, 13 Feb 2017 188 Unknown Fin set Cox’s Bazar

Fig. 3. Molecular species identification using the mitochondrial cytochrome c oxidase subunit I (COI) gene. Neighbour-joining tree of the COI gene (656 bp partial sequence) of specimen 100D, specimen 164, and specimen 44D, and 26 related sequences of Glyphis garricki, G. glyphis, G. gangeticus, and G. fowlerae downloaded from GenBank. Bootstrap values are shown on each branch. The specimens are confirmed as G. gangeticus

3.4. Identification capacity of locals sharks (G. glyphis and G. gangeticus). A total of 87% of the respondents (n = 27) could not differentiate The fishers and traders (n = 31) were asked to any of these species, whereas 13% (n = 4) could tell identify G. gangeticus when shown photographs of that the Ganges shark was a different species from several species from the order Carcharhiniformes. the others. However, they could not specify what the Fishers and traders were asked to differentiate differentiating trait was. The level of identification between G. gangeticus and other species of the order capacity between fishers and traders did not vary Carcharhiniformes. There was no evidence that the significantly. This result is not conclusive with re- fishers and traders (total n = 31) could differentiate spect to their ability to identify the Ganges shark as between leucas and C. amboinensis. the respondents may have been influenced by the They could not differentiate between the 2 river presence of a researcher, biasing their responses to Haque & Das: Ganges shark in Bangladesh 71

these questions. The traders could not elaborate as to reports of G. gangeticus in Bangladesh. The present the exact locations of their catch. However, they study provides clear, diagnostic photographs suitable broadly mentioned nearshore catch locations and for use as future references by researchers and fish- especially the south-central region of Bangladesh ers in Bangladesh. Alternative identification methods and Sundarbans. such as molecular techniques could provide more ac - curate identification for elusive elasmobranch species. Using DNA sequence data, only 3 species of Gly - 4. DISCUSSION phis have been validated (G. gangeticus, G. garricki, and G. glyphis) with one additional species being un- 4.1. The existing G. gangeticus population in described (Li et al. 2015). Hence, molecular tech- Bangladeshi waters niques are recommended for any case of confusion between river shark species until further morpholog- Glyphis gangeticus is one of the most rarely re- ical details are available and incorporated into moni- corded species of shark in Bangladesh, and records toring studies for species confirmation. Altogether, it are extremely patchy throughout its range, with no plays an important role as a case study to evaluate confirmed reports from Bangladesh since 2006. This is the significance of a successful monitoring program due to the rarity of the species and its difficult identifi- based on collaboration with local stakeholders, for cation. Therefore, this publication is the first verified re - better knowledge of the distribution of rare and port of a G. gangeticus specimen from Ban gladesh threatened elasmobranchs for priority conservation within its home range in over a decade. There was actions. only one previous recent record from the Arabian Sea (Jabado et al. 2018). This indicates the importance of studying this elusive species more comprehensively 4.3. Conservation implications of within its range. Although the authors acknowledge this study the limitation of the study due to the small sampling size and the inherent limitations of DNA barcoding in G. gangeticus and G. garricki are assessed as Crit- identifying species, the 3 G. gangeticus specimens, ically Endangered by the IUCN Red List of Threat- one of which was immature, indicate the possible ex- ened Species (Compagno 2007). Even though knowl- istence of a population in the and the edge gaps remain regarding the occurrence, habitat Bay of Bengal. A more comprehensive survey of elas- preference and distribution of G. gangeticus, the mobranch species is therefore critical in this region. present study identifies the importance of systematic surveys in quantifying enigmatic and data deficient species such as Glyphis spp. in survey-poor areas 4.2. Challenges of identification such as the Bay of Bengal. This study additionally highlights the importance of species-specific moni- Identification remains a challenge, as there is a toring and research in Bangla desh and the adjacent lack of photographic evidence to be used as a refer- waters to support evidence-based conservation ence. Historically, one specimen collected from the efforts. Further, it confirms the presence, landing in in 1967 and preserved by the Zoolog- commercial sites, and trade of Ganges sharks in ical Survey of India, Calcutta, was misidentified as Bangladesh and reveals the challenges in identifica- temmnicki ( temminckii), the tion, consistent monitoring, and subsequent lack of . Moreover, Carcharhinus leucas is conservation and research actions. easily confused with G. gangeticus (Martin 2005) and Historical records, museum specimen locality data, is sometimes mislabeled as Carcharhinus sp. due to and the latest studies (Roberts 2006, Amaral et al. their similar morphology (Compagno 2007). This 2017) suggest that the tidal river systems of Sundar- challenge is also demonstrated by the fishers’ and bans and the territorial coastal and marine waters of traders’ lack of ability to identify G. gangeticus and Bangladesh could still be a stronghold of this species. to confuse it with other species. Diagnostic character- There is no systematic marine fisheries monitoring by istics are not practical for use by untrained fishers, the government or fisheries agencies for elasmo- and would lead to potential misidentifications. branch catch in Bangladesh, and all catches are Hence, there is a possibility that the anecdotal bull lumped as ‘sharks, rays and skates’ (FRSS 2018). shark reports from many coastal rivers in Bangladesh Hence, no species-specific data are available from could be G. gangeticus. This could explain the lack of the national statistics, resulting in a lack of historical 72 Endang Species Res 40: 65–73, 2019

understanding of the frequency of landing or bycatch 2005) has a serious potential impact in restricting the trends. Even in the very recent surveys conducted critical habitat for Glyphis spp. within Bangladeshi in India, no specimen was recorded until 2017 (Kiz - waters, while existing trade will further affect the hakudan et al. 2015). This lack of records is indica- population (Haque et al. 2018). More than 69 000 tive of either local extirpation in some of its historical boats deploying more than 183 707 nets are regis- range (Jabado et al. 2018), poor identification capac- tered and in operation (FRSS 2018) alongside a high ities (Akhilesh et al. 2014), or small population size. level of illegal, unreported, and unregulated (IUU) This indicates a data gap in information needed for fishing, all of which are contributing to continued effective conservation measures. elasmobranch bycatch in this area (Shamsuzzaman To collect more species-specific data, a long-term et al. 2017). Due to a possible overlap of the monitoring plan should be implemented and institu- of these exploited marine fish and of G. gangeticus, tionalized within the Bangladesh Fisheries Develop- and the high fishing pressure, it is probable that the ment Corporation, the governing body currently juveniles have been taken as bycatch in the past. responsible for the accounting of marine landing in However, the fishers now retain the catch as there commercial sites. A 3-pronged method of monitoring are markets and demand for these species (Haque et is recommended. al. 2018, A. B. Haque unpubl.). The specimens re ported First, a long-term study at landing sites including in this study were all caught by artisanal fishers and morphological identification using reference photo- all entered into an existing trade of elasmobranch graphs (including teeth), morphometries, and genetic species, almost entirely targeting an international identification is necessary to understand the annual market (Haque et al. 2018). Management measures, landings and bycatch trends and to identify addi- which should urgently prioritize rare and threatened tional species, as suggested by Li et al. (2015). This species as flagship species are therefore mandatory. will resolve the taxonomic challenges posed by these species as well. Second, regular monitoring of the shark processing 5. CONCLUSIONS centres in collaboration with traders is needed to complement the landing site study and achieve a full Given the Critically Endangered status and rarity understanding of catch and trade. of encounters with Glyphis gangeticus, this docu- Third, a thorough socio-ecological study with fish- mentation from Bangladesh is important for species- ers in the coastal waters to understand the catch specific monitoring and management, as well as for trend and identify critical habitats is recommended building a better understanding of its distribution to evaluate any existing populations for evidence- within its range. Understanding the population size based conservation. Environmental DNA (eDNA) in the Bay of Bengal and the adjacent waters and the studies are also recommended to identify the pres- identification of any critical habitats can guide man- ence of elusive elasmobranchs in Sundarbans. agement regimes. It is also important to incorporate The observed specimens of G. gangeticus may the ecological knowledge of fishers into the monitor- have been caught along the coast of the Sundarbans, ing program and to create awareness through in- which would indicate that this is possibly a critical volvement in the management efforts. The authors habitat for the species. The specimens may also have recommend focused research efforts to identify the travelled long distances from the adjacent waters of number of G. gangeticus that are landed annually, Myanmar, India, or Borneo, as marine dis- the presence of any critical habitat and any existing persal of thousands of kilometers has been suggested populations, key sites where trade regulations can be by current gene flow between populations (Li et al. enforced, and a thorough taxonomic study to resolve 2015). As this species uses rivers and coastal areas for all identification issues. nursery grounds and individuals have been recorded from Sundarbans previously (Martin 2005, Roberts Acknowledgements. The authors are grateful to the Bio - 2006, Pillans et al. 2010, White et al. 2015, Feutry et technology Research Centre, the University of Dhaka, for al. 2014), there is a need for transboundary efforts partially funding the genetic work. The authors also thank with a special emphasis on Sundarbans. Moreover, Dr. Peter Kyne and Scott Trageser from the Creative Conser- G. gangeticus is susceptible to the effects of intensifi- vation Alliance for reviewing the manuscript and providing valuable comments, and the International Section Editing cation of anthropogenic modification, habitat degra- Service of the American Fisheries Society for help with edit- dation, and overexploitation (Jabado et al. 2018). ing an earlier version of the manuscript. The authors Imperilment of the Sundarbans (Islam & Wahab acknowledge Hasan Arif Rahman for help with mapping. Haque & Das: Ganges shark in Bangladesh 73

LITERATURE CITED ject, Bangladesh Fisheries Research Institute Islam MS, Wahab MA (2005) A review on the present status Akhilesh KV, Bineesh KK, Gopalakrishnan A, Jena JK, and management of mangrove habitat re - Basheer VS, Pillai NGK (2014) Checklist of chondrichthy- sources in Bangladesh with emphasis on mangrove fish- ans in Indian waters. J Mar Biol Assoc India 56:109−120 eries and aquaculture. Hydrobiologia 542: 165−190 Amaral AR, Smith BD, Mansur RM, Brownell RL, Rosen- Jabado RW, Kyne PM, Nazareth E, Sutaria DN (2018) A rare baum HC (2017) Oceanographic drivers of population contemporary record of the Critically Endangered Ganges differentiation in Indo-Pacific bottlenose (Tursiops adun- shark Glyphis gangeticus. J Fish Biol 92: 1663−1669 cus) and humpback (Sousa spp.) of the northern Kizhakudan SJ, Zacharia PU, Thomas S, Vivekanandan E, Bay of Bengal. Conserv Genet 18: 371−381 Muktha M (2015) Guidance on National Plan of Action for Compagno LJ (1997) Threatened fishes of the world: Glyphis Sharks in India. In: CMFRI Marine Fisheries Policy Series gangeticus (Müller & Henle, 1839) (Carcharhinidae). 2. Indian Council of Agricultural Research, Kochi. www. Environ Biol Fishes 49: 400 fao.org/ipoa-sharks/tools/ipoa-sharks-documents/ en/ Compagno LJV (2001) Sharks of the world: an annotated Li C, Corrigan S, Yang L, Straube N and others (2015) DNA and illustrated catalogue of shark species known to date, capture reveals transoceanic gene flow in endangered Vol 2. Bullhead, mackerel and carpet sharks (Heterdon- river sharks. Proc Natl Acad Sci USA 112:13302−13307 tiformes, Lamniformes and Orectolobiformes). Food and Martin RA (2005) Conservation of freshwater and euryha- Agricultural Organization, Rome line elasmobranchs: a review. J Mar Biol Assoc UK 85: Compagno LJV (2007) Glyphis gangeticus. The IUCN Red 1049−1074 List of Threatened Species: e.T9281A12978210. http:// Moura T, Silva MC, Figueiredo I, Neves A, Muñoz PD, dx.doi.org/10.2305/IUCN.UK.2007.RLTS.T9281A129782 Coelho MM, Gordo LS (2008) Molecular barcoding of 10.en north-east Atlantic deep-water sharks: species identifi- Compagno LJ, White WT, Last PR (2008) Glyphis garricki sp. cation and application to fisheries management and con- nov., a new species of river shark (Carcharhiniformes: servation. Mar Freshw Res 59:214−223 Carcharhinidae) from northern and Papua New Naylor GJ, Caira JN, Jensen K, Rosana KAM, White WT, Guinea, with a redescription of Glyphis glyphis (Müller Last PR (2012) A DNA sequence-based approach to the & Henle, 1839). In: Last PR: White WT, Pogonoski JJ (eds) identification of shark and ray species and its implica- Descriptions of new Australian chondrichthyans. CSIRO tions for global elasmobranch diversity and parasitology. marine and atmospheric research paper 22, CSIRO Bull Am Mus Nat Hist 367: 1−262 Hobart, p 203–225 Pal M, Kar S, Mishra SS (2014) An overview of the fishes of Ebert D, Fowler S, Compagno L (2013) Sharks of the world. Indian Sundarbans and their conservation status. J Envi- Wild Nature Press, Plymouth ron Sociobiol 11: 171−186 Feutry P, Kyne PM, Pillans RD, Chen X, Naylor GJP, Grewe Pillans RD, Stevens JD, Kyne PM, Salini J (2010) Observa- PM (2014) Mitogenomics of the chal- tions on the distribution, biology, short-term movements lenges ten years of control region sequencing. BMC Evol and habitat requirements of river sharks Glyphis spp. in Biol 14: 232 northern Australia. Endang Species Res 10: 321−332 Fischer J, Erikstein K, D’Offay B, Barone M, Guggisberg S Roberts TR (2006) Rediscovery of Glyphis gangeticus: de- (2012) Review of the Implementation of the International bunking the mythology of the supposed ‘Gangetic fresh- Plan of Action for the Conservation and Management of water shark’. Nat Hist Bull Siam Soc 54: 261−278 Sharks. FAO Fisheries and Aquaculture Circular No. Shamsuzzaman MM, Xiangmin X, Ming Y, Tania NJ (2017) 1076. FAO, Rome Towards sustainable development of coastal fisheries re- FRSS (Fisheries Resources Survey System) (2018) Yearbook sources in Bangladesh: an analysis of the legal and insti- of fisheries statistics of Bangladesh, Vol 35. Department tutional framework. Turk J Fish Aquat Sci 17:833−841 of Fisheries, Dhaka Ward RD, Holmes BH (2007) An analysis of nucleotide and Gopal B, Chauhan M (2006) Biodiversity and its conserva- amino acid variability in the barcode region of cyto - tion in the Sundarban mangrove . Aquat Sci chrome c oxidase I (cox1) in fishes. Mol Ecol Notes 7: 68: 338−354 899−907 Haque AB, Biswas A, Latifa GA (2018) Observations of shark Ward RD, Holmes BH, White WT, Last PR (2008) DNA bar- and ray products in the processing centres of Bangladesh, coding Australasian chondrichthyans: results and poten- trade in CITES species and conservation need. Traffic tial uses in conservation. Mar Freshw Res 59: 57−71 Bull 30:6−14 White WT, Appleyard SA, Sabub B, Kyne PM and others Hoq ME, Haroon AY, Hussain MG (eds) (2011) Shark fisheries (2015) Rediscovery of the threatened river sharks, Glyphis in the Bay of Bengal, Bangladesh: status and potentialities. garricki and G. glyphis, in Papua New Guinea. PLOS Support to sustainable management of the BOBLME Pro- ONE 10: e0140075

Editorial responsibility: Austin Gallagher, Submitted: November 2, 2018 ; Accepted: August 10, 2019 Herndon, Virginia, USA Proofs received from author(s): October 5, 2019