Primary productivity connects hilsa fishery in the Bay of Bengal

Abstract

Tropical hilsa shad (Tenualosa ilisha) contributes significantly to the society and economy of Bangladesh, India and Myanmar, but little is known about their habitats across the life cycle and their relationship with environmental drivers. This study describes spatial and temporal variability of productivity in the Bay of Bengal (BoB) relating to hilsa fishery. Decadal data on net primary productivity, nutrients (i.e. nitrate, phosphate and silicate) and zooplankton were collected from Aqua MODIS, world ocean database and COPEPOD respectively with spatial resolution 1°×1°. Moreover, monthly abundance of phytoplankton, hilsa catch and long-term catch dynamics were analyzed to determine the associations between variables. The present study was extended over 3.568 million km2 area, of which 0.131–0.213 million km2 area characterized as the most productive with net primary production of >2,000 mg C/m2/day, 0.373–0.861 million km2 area as moderately productive with 500–2,000 mg C/m2/day, and 2.517–3.040 million km2 area as the least productive with <500 mg C/m2/day which were consistent with field verification data. In case of nutrients, the Ganges-Brahmaputra-Meghna (GBM) delta was rich in nitrate and phosphate than that of the Ayeyarwady delta, while silicate concentration persisted high all over the northern BoB including the deltas. A peak abundance of phytoplankton was observed in GBM delta during the months of August-November, when ~80% of total hilsa are harvested in Bangladesh annually. Variations in seasonal productivity linked with nutrients and phytoplankton abundance are important factors for predicting hilsa habitat and their migration patterns in the deltaic regions and shelf waters of BoB. These results can be useful in forecasting potential responses of the hilsa in BoB ecosystem to changing global ocean productivity.

Introduction

The Bay of Bengal (BoB), which shares many characteristics of the Indian Ocean, is a distinctive system characterized by shallow oceanic arm, deposition of sediment, freshwater plume, seasonal reversal of ocean currents, semidiurnal tides, oxygen-rich surface waters, and abundant biodiversity and fisheries resources1. The northern BoB has the widest shallow shelf region, extending more than 100 nautical miles (=185 km), which is 3–4 times wider in Bangladesh than that in the Myanmar, eastern coast of India and a global average of 65 km. Nutrient is the major driver of primary productivity in BoB2,3. The coupled climate-oceanographic processes (e.g. atmospheric depressions and tropical cyclone, storm surge, internal wave, eddy pumping, and river inputs) are injecting nutrients to the shallow zone and thereby promoting the primary production in upper layers of coastal waters4,5,6, although productivity of an area or ecosystem is dependent on many other bio-physico-chemical parameters such as light7,8.

Phytoplankton is the base of marine food web and thus primary productivity is a key driver of zooplankton and ichthyoplankton dynamics influencing the planktivorous and predatory organisms, mammals, and seabirds9. It accounts for ~50% of global primary productivity10,11, and is important to ecological, biological and biogeochemical processes in oceans12,13. Moreover, marine phytoplankton annually fixes 30–50 billion metric tons of carbon equivalent to 40–50% of the global total14. The majority of phytoplankton species require nitrate, phosphate, iron, molybdenum and copper for growth and reproduction, while diatoms have special requirement of silicate for cell structure and metabolism. Thus, changes in nutrient types and concentrations may change phytoplankton species composition and growth rate, and the resultant net primary production15,16. Zooplankton and herbivore fishes are directly dependent on phytoplankton for food and thus effect primary production through top-down control17. Moreover, productivity (=15–30% of total primary production) is necessary for sustaining the pelagic fish communities18,19. Overall, primary productivity is reported to control the abundance, recruitment, migration pattern and yields of pelagic and migratory fisheries20,21,22, which are important sources of human food, protein, employment and livelihoods17.

The anadromous hilsa shad (Tenualosa ilisha), which feeds on phytoplankton, zooplankton, ichthyoplankton, protozoa, small crustacean and molluscs larvae23,24,25, moves into and out of deltaic ecosystems depending on foraging density and water flow patterns22,26,27. Hilsa acquire energy reserves in the offshore waters until reaching maturity, after which they start upstream spawning migration28, appear to protect reproductive value29,30 with similar behavior in other anadromous fishes, like salmonids and gobiids31,32. Hilsa is mainly harvested from coastal and marine waters (72%), and a quarter from freshwater rivers33. Bangladesh (76%), Myanmar (15%) and India (4%) contribute 95% of the global hilsa catches; while the remaining 5% shared by Iraq, Iran, Kuwait, Thailand and Pakistan34,35. Over the past three decades hilsa production has been increased in Bangladesh with annual harvest reaching 0.5 million tonnes (Fig. 1) that supports livelihoods of 0.5 million fishermen directly, 2.5 million people in the value chain and distribution, and a business valued at US$2 billion33. While peak hilsa fishing takes place from July to November, little is known about their spawning, feeding and/or transboundary migration as well as their spatial and temporal distribution in response to primary production. This study aims to map the productivity zones in the Bay of Bengal and explore their relationship with hilsa fishery.

Figure 1
figure1

Annual hilsa catch in Bangladesh from 1983 to 2018 (left); and few hilsa fishes at a landing center (right).

Materials and Methods

Study area

The present study was carried out in the Bay of Bengal covering an area of 3.6 × 106 km2 between latitude 5° and 24°N, and longitude 79° and 100°E (Fig. 2) covering marine waters of Sri Lanka, India, Bangladesh, Myanmar, Thailand and Indonesia. Major river systems in the study area were Ganges-Brahmaputra-Meghna (GBM) and Ayeyarwady including their tributaries. The BoB has been governed by southwest monsoon winds from May to October and northeast monsoon winds from November to April36,37. Depth of the BoB varies between 10 m in the shelf area of Bangladesh to more than 4500 m as it approaches the Equator38,39. The GBM and Ayeyarwady river systems discharge about 1400 km3/year freshwater, while precipitation (i.e. precipitation over BoB) contributing 5,900 km3/year into the northern BoB40, making it fresher as “river in the sea”41,42. This freshening signal concentrate within the upper 40 m and spreads southward as a narrow strip (~100 km wide) along the western boundary of the bay and reaches the Indian Ocean after 3–5 months journey43,44. Also, freshwater flows southward along the eastern boundary of the basin and gradually mixes with the underlying oceanic layer by vertical exchanges45,46,47. In contrary, the deep canyon in GBM delta (i.e. Swatch of No Ground) has no evidence for sinking freshwater but supports sediment transport to the submarine fan38.

Figure 2
figure2

The Bay of Bengal showing Ganges-Brahmaputra-Meghna (GBM) and the Ayeyarwady deltas (bathymetry in meter).

Data and software

Monthly gridded data during 1998–2018 on net primary productivity (NPP) were collected from Aqua MODIS from NOAA (https://coastwatch.pfeg.noaa.gov) in the area between 5°N–24°N and 79°E–100°E corresponding to the Bay of Bengal. Nutrients data (i.e. silicate, nitrate and phosphate) were collected from the world ocean database (https://www.nodc.noaa.gov/) for the same study domain. Zooplankton data were collected from the Coastal & Oceanic Plankton Ecology, Production, & Observation Database (COPEPOD) (https://www.st.nmfs.noaa.gov/copepod/). The NetCDF format data, which were level 3 products (e.g. SeaWiFS, MODIS and MERIS) with spatial resolution 1° latitude × 1° longitude grid48 (Kodama et al. 2017), were downloaded and geospatially analyzed in ArcGIS software. Marine regions’ data of International Hydrographic Organization (IHO) Sea Areas (version 3) for the Bay of Bengal were collected from http://www.marineregions.org/ and used as base map. The monthly net primary productivity was rated in terms of significance for hilsa habitat modeling (Table 1). The spatial extension module was used for surface interpolation in ArcGIS. All maps and data were transformed into decimal degrees projection. Monthly phytoplankton abundance in the GBM delta was collected from Zafar49. Long term hilsa catch data of 1983–2018 was collected from the Department of Fisheries (DoF), Government of Bangladesh and used for the analysis of hilsa catch dynamics. Besides, monthly hilsa catch data were generated through focus group discussions (FGD) and key informant interviews (KII) with experienced fishers, fish traders and distributors in the coastal fishing villages50.

Table 1 Modeling of net primary productivity.

Data integration

The shape file of study area was converted to grid with cell size of 0.0085 degree, equivalent to 4 km to prepare the mask layer with values 1 inside the mask and 0 outside. The geospatial tabular data of monthly net primary productivity were converted to grids by interpolation, maintaining the same geographic extent and cell size as the mask51. In map calculation, individual parameter grid layers multiplying with the mask layer thus resulted in masked grid layers of corresponding months. These parameter grid layers were then reclassified into three classes with <500, 500–2,000 and > 2,000 mg C/m2/day as most productive, moderately productive and least productive zones (Table 1) respectively, and then evaluated by adding respective months. The two seasons i.e. southwest monsoon (SWM) and northeast monsoon (NEM) were calculated by Eqs. (1 and 2) to develop productivity classification map for hilsa migration in the Bay of Bengal.

$$Gri{d}_{SWM}=({{\rm{Grid}}}_{{\rm{May}}}+{{\rm{Grid}}}_{{\rm{Jun}}}+{{\rm{Grid}}}_{{\rm{Jul}}}+{{\rm{Grid}}}_{{\rm{Aug}}}+{{\rm{Grid}}}_{{\rm{Sep}}}+{{\rm{Grid}}}_{{\rm{Oct}}})/6$$
(1)
$$Gri{d}_{NEM}=({{\rm{Grid}}}_{{\rm{Nov}}}+{{\rm{Grid}}}_{{\rm{Dec}}}+{{\rm{Grid}}}_{{\rm{Jan}}}+{{\rm{Grid}}}_{{\rm{Feb}}}+{{\rm{Grid}}}_{{\rm{Mar}}}+{{\rm{Grid}}}_{{\rm{Apr}}})/6$$
(2)

Validation

Aqua MODIS results of spatial patterns of NPP in the Bay of Bengal were verified after field observation2,6,52,53,54,55,56,57,58,59,60,61. For this, a stratified simple random sampling was used in different habitats (e.g. estuary and delta, shelf water, and high sea) to identify 32 sites for subsequent assessment. Such an approach is appropriate to verify individual locations after the Aqua MODIS have been employed to assess spatial patterns of NPP in the Bay of Bengal. The productivity maps were verified by comparison between predicted productive zones and habitats of hilsa. For this purpose, geographical distribution map of hilsa by FAO, FishBase and GBIF (Global Biodiversity Information Facility), shads distribution by the Shad Foundation, and marine fishing zones of Bangladesh were used. In addition, participatory field visits, 120 semi-structured interviews and 60 focus group discussions were conducted with professional fishers in the coastal fishing villages of Cox’s Bazar, Chittagong, Noakhali, Laxmipur, Bhola and Patuakhali51. In addition, direct observations of hilsa in 55 landing centers were useful and meaningful way to confirm hilsa yields across space and time. The purpose of verification was to find out whether the existing hilsa habitats are in line with productivity classes or not. In addition, members of the academia, researcher and extension manager were consulted to verify the results of geo-spatial models and the distribution of productivity classes of this study.

Results

Net primary productivity (NPP)

The most productive zone is designated with higher NPP and the productivity ranking is presented in Table 2. Monthly and seasonal NPP classification for the study area are illustrated in geo-spatial model (Fig. 3) and summarized in Table 3. After integration of respective months, 0.168 and 0.145 million km2 area were identified as the most productive zone with >2,000 mg C/m2/day during the southwest monsoon (May-Oct) and northeast monsoon (Nov-Apr) seasons respectively. There were 0.636 and 0.489 million km2 moderately productive zone with 500–2,000 mg C/m2/day, and 2.763 and 2.934 million km2 least productive zone with <500 mg C/m2/day (Table 3). The most productive zone was extended over 520 × 90–170 km (lying between 87–92°E and 21–23°N) in GBM delta, and 380 × 40–190 km (lying between 94–97°E and 15–17°N) in the Ayeyarwady delta. Moderately productive zone was confined to 1000 × 30–190 km in GBM delta and 980 × 70–180 km in the Ayeyarwady delta. While, 1540 × 40–130 km along the western boundary (i.e. Indian east coast) and 2130 × 50–180 km along the eastern boundary (i.e. Bangladesh-Myanmar-Thailand coast) of BoB basin were moderately productive. The offshore deep waters and high seas were the least productive zones.

Table 2 Ranking of different habitats according to net primary productivity.
Figure 3
figure3

Monthly and seasonal variations of net primary productivity in the Bay of Bengal.

Table 3 Productivity level and the corresponding area in the Bay of Bengal (total study area 3.568 million km2).

Nutrients

Geo-spatial distribution of nitrate in the upper 10 m indicated that northern and eastern BoB including GBM and the Ayeyarwady deltas were devoid of nitrate (<0.20 μmol/L) during northeast monsoon, but improved situation was evident with >0.60 μmol/L in western BoB along the Indian coast and Sri Lanka during southwest monsoon (Fig. 4). During northeast monsoon the western part of the GBM delta became enrich with nitrate (~0.40 μmol/L) than the east (>0.10 μmol/L), but the Ayeyarwady delta remained with minimum nitrate. The amount of phosphate was >0.4 μmol/L in GBM delta and in western BoB, while higher concentration of >0.6 μmol/L was recorded in the coastal waters (80–120 km wide) of eastern India during southwest monsoon. There were 0.2–0.3 μmol/L phosphate in GBM and Ayeyarwady deltas during northeast monsoon, while lower level (<0.2 μmol/L, a level same as the open ocean) observed in central and southern BoB. Irrespective of season, silicate distribution was higher (>2.5 μmol/L) in northern, western and eastern BoB and also in the deltas (i.e. GBM and Ayeyarwady) possibly due to freshwater influx and residual flow. Lower level of silicate (<2 μmol/L), which is typical for open ocean, was common in southern BoB.

Figure 4
figure4

Spatial distribution of nutrient in the Bay of Bengal.

Zooplankton

Spatial distribution of zooplankton biomass demonstrated that GBM and Ayeyarwady deltas had high zooplankton biomass (> 40 mgC/m3), while reduction of biomass was observed in the western BoB along the Indian coast and eastern BoB along the Myanmar coast (25–40 mgC/m3; Fig. 5). Lower zooplankton biomass (<10 mgC/m3) observed in central and southern BoB. However, the nearshore waters of Sri Lanka, Andaman-Nicobar and Sumatra indicated 15–25 mgC/m3. Spatial distribution of NPP and zooplankton biomass showed that NPP was significantly related with the zooplankton biomass (r = 0.65, p < 0.001). Temporal distribution of zooplankton biomass had minimum variation among the months, i.e. 2.60–2.70 mgC/m3 in January–February, followed by 2.49–2.51 mgC/m3 in June–August and 1.58–1.70 mgC/m3 in October–November (Fig. 6).

Figure 5
figure5

Spatial distribution of zooplankton biomass in the Bay of Bengal.

Figure 6
figure6

Temporal distribution of zooplankton biomass in the Bay of Bengal.

Productivity and hilsa fishery

Concurrent with the nutrient distribution, phytoplankton abundance in GBM delta had an enhanced concentration that also coincided with hilsa catch in the northern BoB at Bangladesh coast (Fig. 7). There were two peaks of phytoplankton in GBM delta of the northern BoB. The first peak of phytoplankton abundance with 3,085 cells/L was recorded in October, while the second peak with 2,470 cells/L was found in March. The peak period of plankton production in August-November was clearly linked to the highest hilsa catch (=~80% of 0.5 million tonnes annual catch), while the second peak in January-March can enhance growth and survival of hilsa juvenile that need scientific investigation. The model outputs for the productivity classes were accurately coincided with available maps of hilsa distribution from various sources such as FAO, FishBase, GBIF, IUCN, Shad Foundation and Discover life (Fig. 8). Acording to the fishermen, estuaries/deltas and nearshore waters are the most suitable zones for hilsa fishing. For that reason they operate hilsa gears within 80 m depth and a distance about 200 km from the coast. This fact suggests that suitable hilsa habitats are distributed in areas where primary productivity is typically high, varifying the model outputs for the productivity classes of this study.

Figure 7
figure7

Phytoplankton abundance and hilsa yields (%) in the Ganges-Brahmaputra-Meghna (GBM) delta of the northern Bay of Bengal.

Figure 8
figure8

Spatial distribution of hilsa in the Bay of Bengal region since 1974.

Among 32 locations (i.e. 4 locations in the estuary and delta, 12 locations in the shelf water and 16 locations in the high sea) selected to verify the patterns of in-situ NPP distribution, the results of 27 locations were comparable to those Aqua MODIS data. Whereas, three locations in the estuary and delta (ID # 1, 2 and 4) and one location in the shelf water (ID # 15) were overestimated, and one location in the shelf water (ID # 9) was underestimated for NPP distribution. Thus, 84.4% of Aqua MODIS output did corroborate with field data (Table 4). The varification error matrix for Aqua MODIS imagery shows the incorrectly classified locations, based on 32 field verification sites (Table 5). MODIS accuracy (MA) and field accuracy (FA) for each of the ranking classes showed that high sea had the highest value of MA and FA (1.0). The shelf water was well discriminated from the rest of the classes (MA = 0.83 and FA = 0.83). The estuary and delta was poorly represented in the sampling (4/32) with MA and FA of 0.05 and 0.25, respectively. The Kappa Index of Agreement (KIA) was generated to determine the degree of agreement between the two outputs. Its value ranges from −1 to +1 after adjustment for chance agreement. A value of 1 indicates that the two outputs are in perfect agreement (no change has occurred), whereas if the two outputs are completely different from one another, then Kappa value is −1. The Kappa (K) and Kendall’s tau (T) coefficients had the value of 0.73 and 0.77 at 95% confidence, indicating that there is good agreement between field reference and Aqua MODIS data for NPP in the Bay of Bengal. We concluded that a high percentage of the pixels was classified correctly, better than would be expected by a completely random classification.

Table 4 Comparison of field observation data against Aqua MODIS ranking of NPP in the Bay of Bengal.
Table 5 Cross-tabulation for error matrix analysis of modeled Aqua MODIS data (row) against field reference data (column) for NPP in the Bay of Bengal.

Discussion

Geo-spatial distribution of nutrients in northern BoB is linked with runoff from GBM and the Ayeyarwady river systems signifying an upward pumping of nutrients from subsurface zone that overlaps with Madhupratap et al.6. Muraleedharan et al.52 recorded <0.1 μM nitrate, 0.4 μM phosphate and >4 μM silicate in the open waters of BoB which is almost similar to present findings. In contrast, Choudhury and Pal53 recorded 11.13–24.19 μM nitrate, 2.23–9.41 μM phosphate and 19.97–127.32 μM silicate along the southeastern coast of India, higher than the present study. Rao et al.62 and De Sousa et al.63 mentioned that rivers flowing into BoB might not contribute much to the inorganic nutrient pool as substantial part of the terrigenous materials are lost at its confluence due to oceanographic processes64. However, river discharges are associated with greater lithogenic fluxes in BoB65, where biogenic matters may rapidly scavenged along with terrigeneous origin and ballasts the materials in faster sedimentation to the deeper ocean66,67. Rivers and atmosphere can supply 20% nitrogenous inputs to BoB, while 80% nitrate comes to the surface from deeper waters by the cyclones, tidal surges, depressions or high speed winds occurring frequently in BoB during Oct-Nov and Mar-Apr68. In general, GBM and the Ayeyarwady deltas are categorized as eutrophic as levels of inorganic nutrients remain high throughout the year compared to the oligotrophic reference values (e.g. phosphate 0.011–0.077 µM, nitrate 0.087–1.900 µM, primary productivity 0.135–0.143 mg Cm−3 h−1) of Karydis69 and Ignatiades70. Continental water flow, nutrients and organic matters originated from the upstream rivers maintain ecosystem functions in the deltas and supply food to the resident species including hilsa71. For instance, the crisscrossed rivers/tributaries of the GBM and Ayeyarwady deltas have an intimate relationship with surrounding land-based activities, such as agriculture, forest, wet meadow, human settlement, industrial development, port operation and tourism activities that play important roles on aquatic habitats. Some areas, such as the offshore deep waters and high seas, are not known as suitable hilsa habitats, but the reasons behind the fact need to uncover with scientific interpretation. Moreover, geo-spatial models can assess suitable habitats of hilsa across the life cycle51, which requires data on bathymetry, oceanographic processes, water quality, primary productivity and habitat characteristics specific to life stages. Some other methods including single nucleotide polymorphism technology (SNP), 87Sr/86Sr isotope ratios in otoliths72, allozymes and morphometric analysis73, and distinctive trait of the parasite fauna74, can provide information about the seasonal movements and residency of hilsa.

High productivity in the nearshore and deltaic waters of BoB integrates with high ambient nutrients. The distribution of NPP positively correlated with phytoplankton concentration (r = 0.75, p < 0.001) that implies that pelagic-neritic fishes like hilsa is expected to maintain their positions within productive areas for sustaining their growth and maturity. Hilsa fed on algae, diatoms, copepods, cladocerans, protozoa, rotifers and the larvae of molluscs24,75. The relationship between hilsa and the abundance of their diets suggests that habitat of hilsa is restricted up to the mixed layer depth (e.g. 20–40 m), which is possibly above the thermocline having maximum level of NPP. At this depth, phytoplankton, copepods, cladocerans and protozoa may be more intense76 and more competently obtained. The present study indicate two peaks of phytoplankton in August-November and in January-March with the lowest abundance in April-July, and this data corresponded well with Choudhury and Pal53 who reported maximum phytoplankton (1,611 cells/mL) in December and minimum (494 cells/mL) in July along the eastern Indian coast of the Bay of Bengal. Incidentally, spawning grounds of hilsa are located in GBM delta77 and Ayeyarwaddy delta78,79, suggesting that these productive water bodies are suitable for hilsa to retain the larvae and juveniles, as illustrated in Fig. 9. In this connection, hilsa migrates to GBM delta for spawning in September-October77 when the levels of NPP and abundance of phytoplankton are high in the system. In addition, coastal rivers and nearshore shallow waters in GBM deltaic region are also suitable nursery grounds for hilsa juveniles until March80. April onwards juveniles start seaward migration as the second phase of anadromous behavior and moves up to 250 km from the coast81 with daily travels of about 71 km82. Moreover, Day83 and Milton84 mentioned that hilsa spends part of life in the sea but not far from the shallow coastal belt. However, operational limitations of gears made difficulties to determine the abundance, extent of seaward migration and fishing potential of hilsa in offshore and high seas due to lack of data and observations. Consequently, Hossain et al.33 recommend for comprehensive study in determining the range of hilsa migration with spatial and temporal distribution across the life cycle. It is evident from the present study that being plankton feeder, hilsa tend to follow plankton rich areas and keep moving from one place to another in search of productive zones and continue to grow. For example, sardine larvae requires 5.7–9.6 mg C/day of primary productivity85 while maximum consumption rates for 1, 2 and 3 years old sardine are 0.042, 0.012 and 0.0049 g-prey g-fish−1 day−1, respectively22. Thus, nursery ground of sardine larvae is located in high productivity zone, where adult sardine (1–3 years old) can live in relatively low productive zone, similar information are not available for hilsa that need to examine. Hilsa prefers zooplankton in early stages and shift towards phytoplankton in adult stage86,87. Diatom, green algae, and blue green algae represent phytoplankton menu of hilsa, while zooplankton menu comprises copepod, cladocera, rotifer, and ostracod23,88,89,90,91. Adult hilsa comprised 97–98% phytoplankton with only 2–3% zooplankton24,92,93 (Fig. 10).

Figure 9
figure9

Conceptual map of hilsa habitats, their migration pattern and life cycle in the northern Bay of Bengal.

Figure 10
figure10

The feeding interactions of hilsa shad.

The relationship between hilsa yield and NPP could be interesting to forecast how hilsa population in BoB might respond to future changes in productivity. The Galathea Expedition in 1950–195254 measured NPP 0.1–0.3 mg C m−2 d−1 in the deep sea and 0.01–2.16 g C m−2 d−1 in the shelf region of BoB6. Data of the subsequent studies from different regions of BoB are given in Table 6. Thaw et al.55 proposed 300–500 g C m−2 year−1 as standard reference value of NPP for eutrophic region. The present study found NPP of >2,000 mg C m−2 day−1 in GBM and the Ayeyarwady deltas that coincided with Thaw et al.55, i.e. 2,590 ± 1,569 mg C m−2 day−1 at the Ayeyarwady delta. Conversely, a drop in primary productivity ranging 500–2,000 mg C m−2 day−1 was noted in the area below 22°N along the western and eastern boundary of BoB basin. The least productivity of <500 mg C m−2 day−1 was found in deeper part of BoB and the Andaman Sea. Though no specific trend was observed in seasonal variations, August-October represented with higher productivity in GBM delta, and lower productivity found in April-June that agrees with the findings of Routray and Patra94, Mohanty et al.13, Kumar et al.2 and Radhakrishna et al.95. Phytoplankton production also follows similar pattern, i.e. phytoplankton abundance is high during August-October in GBM delta. Thus, deltas, estuaries and shelf regions of the northern BoB have higher NPP that may support rich neritic and pelagic fisheries. For instance, Sarmiento et al.95 used coupled climate models to forecast responses of NPP in 2040–2060 and predicted that NPP in the sub-polar North Pacific could increase by 10–20% that could result 20% increase in carrying capacity of pelagic species, such as herring. In this context, distribution of NPP and nutrients in the northern BoB including mega deltas and shelf regions can explain and predict the distribution of pelagic fishes such as hilsa fishery in Bangladesh.

Table 6 Reported net primary productivity (NPP) from different areas/regions of the Bay of Bengal.

Data suggests that total hilsa catch in Bangladesh exceeds nine million tonnes since 1983–84. Specifically, hilsa catch has increased to 517,198 tonnes in 2017–18 from 146,082 tonnes in 1983–84, an annual growth of 7% in the past three decades. The marine waters of northern BoB share the major catch (65%), where the remaining portion fished from the Meghna estuarine system (33%) and from several rivers/tributaries (2%). As the instance in GBM delta, spatial variations in hilsa catch (Fig. 11) interpret habitat suitability and movement routes that can enhance conservation and management initiatives of hilsa fishery. The geographical distribution of hilsa along the coast of Bay of Bengal has been documented by FAO (Fischer and Whitehead 197496; Whitehead97), IUCN98, shad foundation99, FishBase100, Global Biodiversity Information Facility (GBIF)101 and Discover life102 (Fig. 8). Moreover, their occurrences in the adjacent rivers were reported by Hora103, Motwani et al.104 and Quereshi105. Interestingly, hilsa distribution maps with modelled year 2100 native range map based on IPCC A2 emissions scenario (Aquamaps)106 has endorsed similar range for hilsa. Indeed, major efforts of hilsa fishing in Bangladesh have been concentrated within about 100 km from the coast107.

Figure 11
figure11

Spatial variations in catch data explain the habitat and movement routes of hilsa in the Ganges-Brahmaputra-Meghna (GBM) delta (data source: Department of Fisheries, Bangladesh).

The majority of global oceanic data have been collected through cruises for specific periods which is expensive and not many research institutions and country can afford, especially the developing countries like Bangladesh. As a proxy, satellite imagery is useful for specific temporal resolution of any area during the routine observation schedule. For example, MODIS captures data in 36 spectral bands (0.4–14.4 μm wavelength) for 250–1000 m spatial resolutions with viewing swath of 2330 km and the revisit cycle one to two days. MODIS utilizes four on-board calibrators to provide in-flight calibration, whereas vicarious calibration enhances by using marine optical buoy. Thus, the shortwave infrared (SWIR) bands enable more robust atmospheric corrections in turbid coastal waters (Wang et al.)108, while the band around 685 nm is important for detection of phytoplankton fluorescence109,110 (Gower et al., 1999; Hu et al., 2005). Thus, a comparison among the observation/cruises can enhance scientific understanding and interpret the validation process that we applied in this study. Nevertheless, primary productivity is important for pelagic fisheries recruitment, growth and yield, and found to largely controlled by variation in seasonal temperature gradient56, freshwater plume and haline stratification40,43,44,57, vertical transfer of nutrients from the subsurface levels into the euphotic zone111 and dissolved oxygen, which were out of scope of this study. Therefore, further studies are necessary to investigate the multicriteria evaluation of above mentioned parameters to clarify the relationships of hilsa and its habitat conditions.

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Acknowledgements

The Standard NPP datasets is collected from Aqua MODIS of NOAA (https://coastwatch.pfeg.noaa.gov). Nutrients data is collected from the world ocean database (https://www.nodc.noaa.gov/). Zooplankton data were collected from the Coastal & Oceanic Plankton Ecology, Production, & Observation Database (COPEPOD) (https://www.st.nmfs.noaa.gov/copepod/). Marine regions’ data of IHO Sea Areas for the Bay of Bengal is collected from http://www.marineregions.org/. Hilsa fishery catch data is collected from the Department of Fisheries (DoF), Bangladesh (http://www.fisheries.gov.bd) and FAO (http://www.fao.org/fishery/statistics/en). Geographical distribution map of hilsa is collected from FAO (http://www.fao.org/3/ac482e/ac482e00.htm), FishBase (https://www.fishbase.se/summary/1596), the European Nucleotide Archive (EMBL-EBI) Global Biodiversity Information Facility (https://www.gbif.org/species/2413382), Shad Foundation (http://www.cbr.washington.edu/shadfoundation/shad/distribution/), Discover life (https://www.discoverlife.org/20/q?search=Tenualosa+ilisha&b=FB1596) and IPCC A2 emissions scenario Aquamaps (www.aquamaps.org). We are grateful to the NOAA, IHO, DoF, FAO, FishBase, GBIF, Shad Foundation Discover life and Aquamaps for access to these data.

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M.S.H. and S.R.C. conceived and designed the research. S.S. led collection of the data. M.S.H. contributed to data collation. S.S. led data analyses and M.S.H. contributed to data analyses. S.R.C. led illustrations and M.S.H. contributed to illustrations. M.S.H., S.S. and S.M.S. wrote the manuscript. S.R.C. and S.M.S. revised the manuscript. All the authors edited and approved the manuscript.

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Correspondence to M. Shahadat Hossain.

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Hossain, M.S., Sarker, S., Sharifuzzaman, S.M. et al. Primary productivity connects hilsa fishery in the Bay of Bengal. Sci Rep 10, 5659 (2020). https://doi.org/10.1038/s41598-020-62616-5

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