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Copepod community structure during upwelling and non-upwelling seasons in coastal waters off Cochin, southwest coast of India
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Jemi Job N1, *, A. A. Mohamed Hatha1
Acta Oceanologica Sinica | 2019, 38(12) : 111 - 117
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Acta Oceanologica Sinica | 2019, 38(12): 111-117
• Marine Biology •
Copepod community structure during upwelling and non-upwelling seasons in coastal waters off Cochin, southwest coast of India
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Jemi Job N1, *, A. A. Mohamed Hatha1
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  • 1 School of Marine Sciences, Cochin University of Science and Technology, Cochin 682016, India
Published: 2019-12-25 doi: 10.1007/s13131-019-1491-6
Outline
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Vypeen, an island of Cochin estuarine system, acts as the spawning site of several marine and estuarine fishes. We assumed that, physical process (upwelling) make changes in hydrography and the production of chlorophyll a in coastal waters off Vypeen. These alterations can influence the zooplankton abundance and copepod community structure in that area. For justifying this hypothesis, samples were collected from Vypeen at 10 m and 30 m locations during January (pre-southwest monsoon), August (late-southwest monsoon) and November (post-southwest monsoon) 2014. During August, subsurface water column was cool, nutrient rich and less oxygenated (signatures of upwelling) than in November and January. Maximum concentrations of nitrate and chlorophyll a were recorded during August. In the present study, 15 zooplankton groups were recorded; of which copepods were the most predominant group (73%–90%). Copepod density in the present study ranged between 527.2 ind./m3 and 5 139.2 ind./m3. Totally, 37 copepods species were reported in present study and copepod species richness was higher during January. The abundance of zooplankton and copepods was high in August during late-southwest monsoon. These variations were closely associated with the coastal upwelling in August and weakening of moderate upwelling in November. In SIMPER analysis, it was found that the copepods species distribution was similar within season and dissimilar between the three seasons. Moreover, the higher abundance of upwelling indicator species Temora turbinata was recorded in August, which confirms the signs of seasonal upwelling in Vypeen. The present study emphasized on the influence of hydrographical parameters associated with physical process, in governing the copepod community organization of the Vypeen Island.

copepods  /  community  /  multivariate analysis  /  monsoon  /  Temora turbinata  /  upwelling  /  zooplankton
Jemi Job N, A. A. Mohamed Hatha. Copepod community structure during upwelling and non-upwelling seasons in coastal waters off Cochin, southwest coast of India[J]. Acta Oceanologica Sinica, 2019 , 38 (12) : 111 -117 . DOI: 10.1007/s13131-019-1491-6
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1 Introduction
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Copepods are the dominant mesozooplankton in the marine environment, especially in the estuaries and coastal regions where they contribute over 80%–90% (Srichandan et al., 2014) of the total zooplankton abundance. Most of the recent literatures that reported on the zooplankton and copepods in the Indian waters with respect to the ocean physics are related to the upwelling (Fernandes and Ramaiah, 2009; Rakhesh et al., 2008, 2006). However, recent literature on the zooplankton and copepod species diversity from Kerala coast (Cleetus et al., 2016; Vineetha et al., 2015; Jeyaraj et al., 2014; Robin et al., 2009) is very limited. Similarly, comparative studies of the copepod community structure during upwelling period with respect to the non-upwelling periods along Southwest coast of India and especially from Kerala are meager (Jagadeesan et al., 2017; Sooria et al., 2015; Vineetha et al., 2015; Varghese and Krishnan, 2009; Madhupratap, 1978; Tranter and Abraham, 1971).
In the southwest coast of India the reversal of the coastal currents and prominent upwelling during the southwest monsoon period, brings dynamic changes (Bhattathiri et al., 1996) in estuaries and coastal waters. This region is also experienced by the winter cooling and the maintenance of weakened upwelling during the northeast monsoon (Smitha et al., 2014). Also, the seasonally varying river flow make changes in hydrography and chlorophyll a levels along the Cochin estuary (Shivaprasad et al., 2013). Vypeen is considered as a highly bioproductive coastal area of Cochin Backwaters along southwest coast of India. Copepods were found to be the predominant group among zooplanktons in present study. Considering this importance, the present study was carried out to evaluate the zooplankton distribution and copepod species diversity related to hydrography during upwelling and non-upwelling periods in Vypeen.
2 Materials and methods
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2.1 Study area
The study area was distributed along the latitudes 9°N and 10°N (southwest coast of India). The Vypeen Island (10 m location–latitude 9.952 83°N and longitude 76.286 1°E; 30 m location–latitude 9.952 78°N and longitude 76.188 8°E) is the transect which selected for the study. This transect is located in Kerala along southwest coast of India. Two locations in Vypeen transect were selected for sampling, one is at 10 m and another one is at 30 m (Fig. 1). A mixed, semidiurnal type tides with a maximum range of about 1 m usually experiences along this coastal area off Cochin Backwaters. Vypeen is a highly bioproductive coastal area of southwest coast of India which is bound by the Arabian Sea on western side and the Cochin Backwaters on eastern side.
2.2 Sampling and methods
Samples were collected in three seasons from Vypeen at 10 m and 30 m locations. For the comparison of seasonal changes, sampling was carried out in January 2014 (pre-southwest monsoon), August 2014 (late-southwest monsoon) and November 2014 (post-southwest monsoon). At each location, water samples were collected from the surface (0.5 m) using Niskin sampler (General Oceanics, USA) for the estimation of chlorophyll a, nutrients and dissolved oxygen. The vertical distribution of physical parameters such as salinity and temperature were recorded by factory calibrated sensors. Temperature was measured by using a thermometer and salinity by using a calibrated probe, immediately after taking the water sample at each station. The dissolved oxygen in the surface and subsurface waters was measured by Winkler’s method (Grasshoff et al., 1983). The total chlorophyll a was measured by concentrating 1 L of water samples on Whatman GF/F filters (0.2 μm) and was extracted using 10 mL 90% acetone. The extracted chlorophyll a was measured using a Trilogy Turner fluorometer following the standard procedure (UNESCO, 1994). Nitrate concentrations in water were measured by following Grasshoff’s method (Grasshoff et al., 1983). Zooplankton samples were collected by horizontal hauls using WP-2 net (200 µm mesh size), attached with a calibrated digital flow meter (Hydro-bios, Germany). After each haul the zooplankton samples were transferred into cleaned 0.5 L plastic containers and were fixed in (4%–5%) buffered formalin. The samples were filtered through a 200 µm nylon sieve in the laboratory and excess water in the samples was removed using blotting papers.
Zooplankton samples were evaluated qualitatively and quantitatively in the laboratory by following the standard procedures of Postel et al. (2000). The zooplankton sample was split into four sub-samples (25%) using a Folsom’s plankton sample splitter (Sell and Evans, 1982). One of these sub-samples (25%) was sorted under a stereozoom microscope (Model–Mastic DMW–143–FBGG) and taxonomic group level abundance was estimated (Postel et al., 2000). Among various taxonomic groups, copepods were analyzed thoroughly and their species were identified using standard keys (Kasturirangan, 1963; Conway et al., 2003). The abundance of copepods was expressed as ind./m3. Zooplankton diversity indices-species diversity index (H), species richness (d), evenness index (J′) were evaluated by applying the formulas of Shannon and Weaver (1963), Margalef (1968) and Pielou (1969) respectively.
2.3 Statistical analysis
In order to compare the hydrography and the zooplankton characteristics in Vypeen, univariate and multivariate analysis was done. Based on biological parameters, sampling locations were segregated using hierarchical agglomerative method of cluster analysis. The data of copepods species abundance were initially log10(X+1) transformed for normalizing the differences in numerical abundance (Clarke and Warwick, 2001). The Bray-Curtis similarity matrix was used for the spatial grouping of locations in different seasonal collections. In addition to cluster analysis, similarity profile (SIMPROF) permutation test was also performed to identify significant assemblages of stations (p<0.01) (Clarke and Gorley, 2006). The relationship between copepod species and some essential environmental variables were evaluated by using RDA (CANOCO 4.5). Samples were displayed by points and species and quantitative environmental variables were shown by arrow in RDA triplots.
3 Results and discussion
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3.1 Hydrography and chlorophyll a
The temporal distribution of surface temperature showed 23.4°C to 31.5°C variations between January, August and November (Table 1). During January the water column was little warmer (31.5°C and 30.9°C at 10 m and 30 m locations respectively), than the months of August and November. Surface salinity ranged from 31.06 to 34.63 in present study (Table 1). Surface salinity was comparatively less during August, but remarkably subsurface salinity was high compared to January. The high surface water temperature and salinity during the pre-southwest monsoon period (January) along the Arabian Sea has been reported by Madhupratap (1978). During the southwest monsoon period salinity values in coastal waters changes due to the freshwater inputs from the outlets of Cochin Backwaters. In present study, the sea surface temperature and salinity were less in the Vypeen during the late-southwest monsoon period. Similar changes about the salinity and temperature has been reported by Haugen et al. (2002) and Madhupratap (1978) previously from Cochin Backwaters. During the northeast monsoon (November) period the coastal water salinity becomes intermediate between late-southwest monsoon period (August) and pre-southwest monsoon period (January). These observations were accordance with the earlier observations like Madhupratap (1978), Madhupratap et al. (2001) from the South Eastern Arabian Sea.
In general nitrate concentration was two to five folds higher during August compared to January. During August surface nitrate concentrations were 5.69 µmol/L at 10 m and 4.91 µmol/L at 30 m locations (Table 1). The lowest concentration of nitrate was found during January compared to August and November in the present study. The previous investigation by Rajagopalan et al. (1992) has recorded low surface water nitrate concentration during pre-monsoon along southwest coast of India. The highest concentration of dissolved oxygen in surface waters was recorded during August (6.72 mg/L) and the lowest concentration of dissolved oxygen in subsurface waters was found in the same season (August). It indicates the subsurface hypoxia during upwelling period (Table 1). The higher nitrate concentrations, reduced temperature and less oxygenated water in the Arabian Sea may be the result of the coastal upwelling and winter cooling during southwest monsoon, northeast monsoon periods respectively (Gerson et al., 2014).
Generally chlorophyll rich waters were observed in upwelling areas by Bhattathiri et al. (1996) and Rajagopalan et al. (1992). Chlorophyll a is regarded as the indirect representation of the phytoplankton production. Chlorophyll a concentrations ranged from 0.40 mg/m3 to 8.96 mg/m3. The concentration of chlorophyll a was very less during January which denotes the low productivity off Vypeen, in the pre-southwest monsoon period during 2014. The maximum chlorophyll a concentration was found during August and moderate during November in the present study (Table 1). These observations confirm the upwelling signatures during August (Late-Southwest Monsoon) and the winter cooling due to the weakening of moderate upwelling during November (post-southwest monsoon) in Vypeen.
3.2 Zooplankton distribution and copepod diversity
Zooplankton abundances showed significant seasonal variation in present study. A total of 15 zooplankton groups were reported in the present study (Table 2). Density of the zooplankton varied from 660.6 ind./m3 to 7 001.9 ind./m3. Zooplankton abundances were higher during August and November, but it was found to be lower during January in Vypeen. Copepod contribution to the total density was high (>80%) in 10 m location compared to the 30 m location. The maximum density of copepod (90%) was found during the late-southwest monsoon period (month of August at 10 m location). Cladocerans contributed (0.9%–23.2%) to the total density of zooplankton; higher density was observed during August and November compared to January. The copepod predators like Chaetognaths, Siphonophores and Hydromedusae density was high during August and November compared to January. The other holoplankton groups found in the total density was Lucifer, Mysids, Thalacians, Appendicularians, but they contributed 1%–2% to the total density of the zooplankton. Three different groups of the meroplankton was found such as polycheate larvae, molluscan larvae, decapod larvae their contribution was found 0.2%–5% to the total density. Variations in abundance of zooplankton and copepods in the Cochin Backwaters during upwelling and non-upwelling seasons have been recorded by Sooria et al. (2015), Vineetha et al. (2015) and Varghese and Krishnan (2009). In the present study, the most abundant group was copepod (≥80%). The previous studies from Cochin Backwaters showed that copepods were the most predominant group among zooplankton (Vineetha et al., 2015; Madhupratap, 1979). Recently, higher abundance of mesozooplankton, especially the dominance of copepods, along the southwest coast of India (Alappuzha in Kerala) during the southwest monsoon period governed by coastal upwelling has documented by Jagadeesan et al. (2017).
Copepod density in the present study ranged between 525.2 ind./m3 and 5 139.2 ind./m3 (Table 2). Copepod density was high in 30 m location during August and November compared to the 10 m location. But the differences between 10 m and 30 m locations were comparable. In the present study, altogether 37 species were reported (Table 3). Among these, 21 species belonged to the order Calanoida, 2 species belonged to the order Harpacticoida, 5 species belonged to Cyclopoida order, 8 species belonged to the order Poecilostomatoida and 1 species belonged to the order Monstrilloida. In Calanoida order, 9 families were found such as Calanidae, Eucalanidae, Paracalanidae, Centropagidae, Pseudodiaptomidae, Temoridae, Candaciidae, Pontellidae and Acartiidae. In Harpacticoida order 2 families were found such as Macrosetellidae, Tachidiidae.
Copepod species distribution pattern during August (upwelling period), November (moderate upwelling) and January (non-upwelling period) showed variations. Especially, the abundance of Temora turbinata, Oithona similis, Centropages orsini, Paracalanus parvus, Centropages tenuiremis, Corycaeus catus and Acartia erythraea was significantly varied. The abundance of the Temora turbinata (upwelling indicator species) was found several folds higher during the late-southwest monsoon period (August) compared to the pre-southwest monsoon and post-southwest monsoon periods (January and November) in the Vypeen transect. Similar observations has been reported during upwelling season along Indian coastal waters by Jagadeesan et al. (2017, 2013), Goswami and Padmavati (1996) and Madhupratap et al. (1990). Copepod species richness was high during the month of January and the minimum species richness was reported during the month of August. Shannon diversity of copepods was high in January (10 m location) compared to the months of August and November. Pielous evenness showed noticeable variations between the observations. Maximum evenness was found during the month of August in 30 m location but during the month of January the evenness values were comparable in 10 m and 30 m locations.
3.3 Cluster analysis and NMDS
Spatial and temporal similarity of species distribution of the observations from the Vypeen transect (January, August, November 2014) were grouped into three clusters at the similarity level of 65% (Fig. 2a). Cluster 1 included the 10 m and 30 m locations of the month of January. Second cluster included the 10 m and 30 m locations of the month of August and the cluster 3 included the 10 m and 30 m locations of the month of November. It indicated that the classification of the clustering pattern was significant and the species distribution was significantly differed between the clusters. Cluster overlaid NMDS plot shows that 10 m and 30 m locations of the each month was located close to each other represents the similarity (Fig. 2b). But the points which are located far between the months represent the dissimilarity.
3.4 SIMPER analysis
SIMPER analysis (similarity percentage) was performed to find out the species distribution similarity within the cluster and between the clusters. Similarity percentage analysis also represented the dissimilarity of species distribution and provided the information about the dominant species responsible for the dissimilarity. The results of the SIMPER analysis were represented in the Supplementary Information Table S1. Table S1a, b and c provide the information about similarity percentage within the cluster members in Group 1, Group 2 and Group 3, respectively. Table S1d, e and f presents the information about the dissimilarity between Groups 1 and 2, Groups 1 and 3, and Groups 2 and 3 members, respectively. Cluster 1 group members (the locations of 10 m and 30 m during January) showed 61.30% similarity within the cluster. In Cluster 2 (10 m and 30 m locations, during August) the species distribution showed 57.54% similarity within the cluster members. Cluster 3 members (10 m and 30 m locations of November) contributed 60.79% similarity within cluster. The average dissimilarity between Cluster 1 and Cluster 2 was 88.57%. The average dissimilarity between Cluster 1 and Cluster 3 was 91.91%. The average dissimilarity between Cluster 2 and Cluster 3 was 78.99%.
3.5 Redundancy analysis (RDA)
RDA was carried out to analyze the relationship between the copepod species and the environmental variables in Vypeen during the pre-southwest monsoon, late-southwest monsoon and post-southwest monsoon periods in 2014 (Fig. 3). Salinity and temperature oriented the same direction indicated that both were positive relationship to each other. The nitrate was oriented opposite to the direction of the temperature and salinity in RDA triplot that indicated its inverse relationship to the temperature and salinity. The zooplankton density, surface and bottom chlorophyll a and the species of Acartia danae, Acartia erythraea, Oithona similis, Centropages tenuiremis, Temora turbinata, Temora discaudata, Centropages orsini and Paracalanus parvus oriented to the right hand direction and that showed the positive relationship to one another.
Generally, Oithona similis occur in the subsurface waters and their abundance shall be less in the surface waters. However, high abundance of Oithona similis was found in the surface waters during August (late-southwest monsoon) and November (post-southwest monsoon) respectively in the present study. The similar observation was reported from estuarine waters of Washington by Keister and Tuttle (2013). Their investigations suggested that the Oithona may be migrated to the surface layers due to subsurface hypoxia during upwelling seasons. Lowest species richness and highest evenness values were found in the late-southwest monsoon period compared to non-upwelling periods. Similar observation of decreasing copepod species diversity, when one or two species dominated to the total density, during southwest monsoon period was reported from upwelling area along the central west coast of India (Madhupratap et al., 1990).
4 Conclusions
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Results of community structure of copepods in coastal waters off Vypeen revealed that distribution and abundance of the copepod species were differed between the seasonal observations associated with physical processes. During August, water column was cool, nutrient rich and less oxygenated (signatures of upwelling) and high in chlorophyll a than in November and January. The abundance of zooplankton and copepods was high in August during late-southwest monsoon. High abundance of Temora turbinata reported in the study, confirms upwelling during southwest monsoon. These variations were closely associated with the coastal upwelling in August and weakening of moderate upwelling in November.
Acknowledgements
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We thank the scientist–in–charge, CSIR-National Institute of Oceanography, Regional Centre, Kochi, India for facilities and encouragement. We express our sincere thanks to R. Jyothibabu, from CSIR-NIO, Regional Center, Kochi for valuable suggestions and support. We thankfully acknowledge Head of the Department of Marine Biology, Microbiology and Biochemistry, Cochin University of Science and Technology, Fine arts Avenue, Cochin, India for providing the facilities. We also thank Ministry of Earth Science (MOES), Government of India for financial support through ITIS programme.
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Year 2019 volume 38 Issue 12
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doi: 10.1007/s13131-019-1491-6
  • Receive Date：2018-05-04
  • Online Date：2026-04-01
  • Published：2019-12-25
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  • Received：2018-05-04
  • Accepted：2018-11-15
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    1 School of Marine Sciences, Cochin University of Science and Technology, Cochin 682016, India

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表12种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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