Benthic foraminiferal responses to bottom water characteristics in the palk bay, off Rameswaram, southeast coast of India. Study revealed that the microfaunal assemblage consisting of 108 species belonging to 50 genera of 27 families. Increase in temperature, salinity and dissolved oxygen content of the bottom water were observed as favourable factors promoting the abundance of living population.
Benthic foraminiferal responses to bottom water characteristics in the palk bay, off Rameswaram, southeast coast of India. Study revealed that the microfaunal assemblage consisting of 108 species belonging to 50 genera of 27 families. Increase in temperature, salinity and dissolved oxygen content of the bottom water were observed as favourable factors promoting the abundance of living population.
Benthic foraminiferal responses to bottom water characteristics in the palk bay, off Rameswaram, southeast coast of India. Study revealed that the microfaunal assemblage consisting of 108 species belonging to 50 genera of 27 families. Increase in temperature, salinity and dissolved oxygen content of the bottom water were observed as favourable factors promoting the abundance of living population.
Benthic foraminiferal responses to bottom water characteristics in the Palk Bay, off Rameswaram, southeast coast of India
1 V Kumar & 2 V Manivannan 1 Department of Geology, National College, Tiruchirapalli-620 001, Tamil Nadu, India 2 Department of Geology, Government College, Salem 636 007, Tamil Nadu, India Received 29 August 2000, revised 10 May 2001 Recent benthic foraminiferal assemblages from the shallow shelf sediments of the Palk Bay, off Rameswaram were studied qualitatively and quantitatively. The study revealed that the microfaunal assemblage consisting of 108 benthic fo- raminiferal species belonging to 50 genera of 27 families. Various bottom water parameters were determined and correlated with the foraminiferal population and seasonal distribution of living population was evaluated. The ecology and distribution of eight widespread and abundant foraminiferal species of the study area viz. Spiroloculina communis, Quinqueloculina seminulum, Triloculina insignis, T. trigonula, Ammonia beccarii, A. tepida, Pararotalia nipponica and Osangularia venusta were discussed in detail. Increase in temperature, salinity and dissolved oxygen content of the bottom water were observed as favourable factors promoting the abundance of living population in the study area. Foraminifera occur in marine and brackish water en- vironments. The occurrence of benthic species is con- trolled to a great extent by physical factors viz. depth, temperature, amount of light, turbidity of the water; chemical factors of water viz., salinity, dissolved oxygen and available elements; biological factors viz., available food supply and character of the bottom sediments 1-3 . Studies have been conducted on the ecology and distribution of Recent benthic foraminif- era from east coast 4-10 and from west coast 11-15 of In- dia. Naidu et al. 6 reported, after studying the fo- raminifera from Visakhapatnam harbour complex, that foraminifera can be used as an indicator for ma- rine pollution. Sreenivasa Rao et al. 8 studied the dis- tribution of foraminifera from Nizampatnam bay and determined four assemblages which are influenced by distinct environmental conditions prevailed in the area by means of Q-mode factor analysis. Khare et al. 15 studied the foraminifera from off Mangalore and concluded that the distribution pattern of benthic foraminiferal morpho-groups can be correlated with fresh water river discharge. The present work was undertaken to study the influence of bottom water characters on the foraminiferal population and diver- sity in general, and ecology and distribution of abun- dant and widespread foraminifera, in particular. Materials and Methods Fifty-two sediment and bottom water samples were collected at 13 sampling stations, once in three months, representing four seasons in the year 1987, from the Palk Bay, off Rameswaram (long. 79 29' to 79 35' E and lat. 9 15' to 9 17'N) (Fig. 1). The sam- ples were collected in depths ranging from <1 m to 16 m. Unit volume of 25 ml wet sediment from each sample was preserved in formaldehyde solution and the living and dead populations of the foraminifera were quantitatively and qualitatively examined for the ecological studies 16 .
Fig. 1Map showing location of sampling stations in the Palk Bay, off Rameswaram, southeast coast of India INDIAN J . MAR. SCI., VOL. 30, SEPTEMBER 2001
174 The temperature of the bottom water sample was recorded from the built-in thermometer. A portion of water sample at each station was preserved according to Strickland & Parsons 17 to estimate the dissolved oxygen content. To determine the salinity of water, chlorinity was estimated and related by Knudsen equation. The pH values were measured for each of the water sample using pH meter. Nutrients (silicate, phosphate, and nitrate) were estimated using methods described by Strickland & Parsons 17 .
Results and Discussion
Bottom water Temperature: In the present area, since the samples were collected from shallow depths, there was no ap- preciable variation in temperature among the different stations of a season (Figs 2 and 3). However, there was an appreciable variation in temperature during different seasons (Fig. 4). It was generally observed that the population of living foraminifera was more when the temperature was high. Salinity: In the bottom water, off Rameswaram, the salinity values (Figs 2 and 3) varied from 30 (stations 6, 7 -J anuary) to 35.2 (station 5,7 - J uly). The sa- linity values did not show any noticeable variations within the stations in particular season, but there was a conspicuous variation among different seasons. The average salinity value was higher during J uly, which was very closely followed by April (Fig. 4). In the study area, it was observed that the living population of foraminifera was more during these two seasons, which may be attributed, mainly, to salinity values ranges from 33.6 to 35.2 . Dissolved oxygen: The distribution of dissolved oxygen showed a clear seasonal variation in the
Fig. 2Showing the relation between the cumulative percent of widespread and abundant living foraminifera and bottom water parameters (J anuary and April) KUMAR & MANIVANNAN: BENTHIC FORAMINIFERAL RESPONSES
175 bottom waters, off Rameswaram (Figs 2-4). The maxi- mum mean value was observed in the month of April and the minimum in J anuary. Spatially, there was no observable variation noticed among the different sta- tions of a season. The increase in the dissolved oxy- gen content during April may be due to high photo- synthetic activities of phytoplankton 1,4 . This increase in the dissolved oxygen content coincided with the increase in the living population in the present area, as reported for Vellar estuary by Ramanathan 19 . It was considered that the dissolved oxygen content was one of the important factors governing the standing crop. pH: According to Phleger 20 hydrogen ion concen- tration is expected to affect the production of calcare- ous tests so that at a pH approximately <7, such forms may not survive. In the present study, the pH values of the bottom water did not show much varia- tion spatially and seasonally (Figs 2-4). Since the fluctuation in the pH values was meager, the water maintained alkalinity throughout, within the limits of the tolerance of the fauna. Because of the small varia- tions in pH, its effect on the distribution of foramini- fera was not possible of accurate determination. Nutrients: The variations in the concentration of nutrients-silicate, phosphate and nitrate constituents of the bottom waters for the different stations of dif- ferent seasons are given in Figs 2 and 3. As the distri- bution of all the nutrients was meager and erratic spa- tially, the correlation between them and the living crop cannot be achieved. But it was observed from the mean values (Fig. 4) that the increase of these nutri- ents showed an inverse relationship with the popula- tion. Foraminifera The qualitative analyses of sediment samples led to the recognition of 108 benthic species consisting of
Fig. 3Showing the relation between the cumulative percent of widespread and abundant living foraminifera and bottom water parameters (J uly and October) INDIAN J . MAR. SCI., VOL. 30, SEPTEMBER 2001
176 both living and dead fauna. They belong to 50 genera, 27 families and 10 superfamilies 18 . Among the 108 species, 12 species [viz. Rhabdommina scabra, Ammobaculites exiguus, Textularia agglutinans, T. aura, T. candeiana, T. conica, T. foliacea, T. foli- acea occidentalis, T. palustris,. Bigenerina irregu- laris, Trochammina inflata, and Eggerella advena] are arenaceous agglutinated (suborder Textulariina), 43 are calcareous porcelaneous [viz. Edentostomina cul trata, Spiroloculina angulata, S. communis, S. corru- gata, S. costifera, Spiroloculina. sp., Vertibralina striata, Quinqueloculina agglutinans, Q. bicostata, Q. bidenta compressa, Q. lamarckiana, Q. parkeri, Q. polygona, Q. pseudoreticulata, Q. rameswarensis, Q. seminulam, Q. sulcata, Q. tenagos, Q. undulose costata, Pseudomassilina australis, P. australis re- ticulata, P. macilenta, Pyrgo elongata, P. sub- spherica, Triloculina carinata striata, T. insignis, T. schreiberiana, T. terquemiana T. tricarinata T. trigonula, Rupertianella rupertiana, Miliolinella circularis, M. labiosa, Hauerina bradyi, H. fragilis- sima, Articulina mayori, Parrina bradyi, Peneroplis plantus, Monalysidium politum, Spirolina acicularis, S. arietinus, Sorites marginalis and S. orbiculus (Sub- order Miliolina)] and the rest 53 are calcareous perfo- rate [viz. Lagena costata amphora, L. gracillima, L. laevis, L. setigara, L. striata, Guttulina sp,. Fissurina marginata, Bolivina doniezi, B. lanceolata, Brizalina lowmani, B. striatula, Rectobolivina glabra, R. raphanus, Chrysolidinella dimorpha, Reusella at- lantica, Rosalina globularis, R. valvulata granulosa, Cancris oblonga, Spirillina vivipara, Ammonia bec- carii, A. dentata, A. tepida, Asterorotalia inflata, A. trispinosa, Pararotalia nipponica, P. ozawai, Pseudorotalia schroeteriana Calcarina umblicata, Elphidium advenam, E. crispum, E. discoidale, E. excavatum, E. hispidium, E. incertum, E. limbatum, E. poeyanum, E. verriculatum, Poroeponoides later- alis, Cibicides lobatulus, C. refulgens, Planorbulina mediterranensis, Planorbulinella larvata, Acervulina inhaerens, Cymbaloporetta bradyi, C. squammosa, Fursenkoina compressa, F. punctata, Sigmavirgulina tartuosa, L. limbatum costulata, Florilus boueanum, F. grateloupi, F. labradoricum and O. venusta (sub- order Rotaliina)]. Out of the 52 sediment samples collected and ana- lysed from the Palk Bay, the living foraminifera have been observed in all the samples (Table 1). The living population was maximum during April (summer sea- son) and abundant in the intermediate stations (sta- tions 4 to 9) irrespective of any season. Among the 108 species identified, only 85 species were found to be in living condition, which include 9 arenaceous, 34 porcelaneous and the rest 42 were perforate forms. Among living, only the eight taxa [viz. Spiroloculina communis, Quinqueloculina seminulum, Triloculina insignis, T. trigonula, Ammonia beccarii, A. tepida, Pararotalia nipponica and Osangularia venusta] were considered to be widespread and abundant in the pre- sent area since they were found in living condition in more than 75% of the samples collected and studied.
Distribution of foraminifera Out of the 108 benthic foraminiferal species recog- nised in the Palk Bay, 23 were not found in living condition and such species are collectively referred as dead species in the present study. Among the rest 85 which are found living, 9 are arenaceous aggluti- nated, 34 are calcareous porcelaneous and the rest 42 are calcareous hyaline. The following 8 species are considered to be widespread and abundantly occur-
Fig. 4Seasonal mean values of living population and bottom water parameters KUMAR & MANIVANNAN: BENTHIC FORAMINIFERAL RESPONSES
177 ring since they are found as living in more than 75% of the samples collected and studied: Spiroloculina communis, Quinqueloculina seminulum, Triloculina insignis, T. trigonula, Ammonia beccarii, A. tepida, Pararotalia nipponica and Osangularia venusta. The foraminiferal population (living) was found to be more in the summer season (April). This abun- dance of population may be positively correlated with an increase of temperature, salinity and dissolved oxygen content of the bottom waters. Since there was no observable variation of these bottom water charac- ters among different stations of a season, a positive correlation between them and population cannot be achieved. Kumar et al. 9 have reported that a silty sand substrate and higher calcium carbonate content of sediments favoured an abundant population. The rela- tion between the cumulative percent of abundant and widespread living foraminifera and the various bot- tom water parameters governing them are given in the Figs 2 and 3. Spiroloculina communis Cushman & ToddThe living representatives were absent only in the shore stations of April and October. Spatially, this species was found to be abundant in the later half stations (7 to 13) of the traverse. As there was no much variation in the bottom water parameters among the stations, it was hard to correlate them with the population of the species. Temporally, this species had higher popula- tion in summer and lesser during October. This popu- lation variation can be very well correlated positively with the increasing temperature, dissolved oxygen content of the bottom water. The optimum salinity value for higher reproduction of this species is be- tween 33.6 and 34.8. Quinqueloculina seminulum (Linn)The living specimens of Q. seminulum occurred in 41 samples. Spatially, more population was noticed in the middle segment of the traverse (stations 3 to 9). The living forms were found to be abundant during April and minimum during October. The temporal distribution of this species can be positively correlated with the higher temperature and dissolved oxygen content of the bottom water. The optimum salinity value for higher reproduction was <3. Kumar et al. 9 have observed that the higher calcium carbonate content of the substrate was the main controlling factor for the abundance in the intermediate segment of the traverse. Triloculina insignis (Brady)The living speci- mens were absent in the first and last stations of all the collections. In general, the living specimens were more concentrated in the intermediate stations (4 to 10). Temporal distribution of the species revealed that the living population was greater during summer (April) followed by J anuary. During the monsoon seasons the living population size was considerably small. In this area, it was found that the active repro- duction for T. insignis occurred between J anuary and April, with more reproductive activity in April. Triloculina trigonula (Lamarck)The living population was completely absent in the shore sta- tion and minimal in the second. The concentrations were more in the intermediate stations of the trav- erse. Temporally, the maximum living population was seen in summer (April) which was followed by winter (J anuary) while both the monsoons witnessed a lower living population size. Complete absence of living forms in the first station as well as the mini- mum in the second station showed that the near- shore turbulence caused by the wave action was not favourable for the thriving of T. trigonula. The smaller population size during J uly and October may be due to the monsoon weather and bay depressions. This species prefers quiet water as in the case of T. insignis. Ammonia beccarii (Linne)Phleger & Lankford 21
reported that the reproduction of Streblus beccarii (Linne) occurs in frequent intervals and is not related to the season. Boltovskoy 22 stated that Rotalia bec- carii did not show any seasonal periodicity and as- sumed that reproduction occurred at frequent intervals throughout the year. The spatial distribution of this species in the present area was erratic. Seasonally, the Table 1Living foraminiferal populations, off Rameswaram (No. of specimens/25 ml. of wet sediment)
178 living population size was maximum during SW mon- soon followed by winter and minimum during NW monsoon and summer. Ammonia tepida (Cushman)Spatially, the distri- bution of A. tepida reveals that they were compara- tively lesser in the last 3 stations of any traverse. Temporally, the total living population for a season was found to be maximum during April, followed by J anuary and was minimum during October. Brad- shaw 23 observed from the laboratory experiments that the reproduction of A. tepida occurred only between 20 - 30C, with optimum conditions from 25 - 30C and salinity values between 20 40. In the present study, this species occurs in living condition in almost all the samples, indicating its tolerance to consider- able upward and downward fluctuations of environ- mental factors. The distribution of the fauna of the present area reveals that the optimum conditions for the abundance of this species aretemperature 31.5C - 32.6C; dissolved oxygen - 6.0 ml/l; salinity <34.5and low nutrient values. Pararotalia nipponica (Asano)The distribution of P. nipponica reveals that the population was found to be more in stations between 1 and 7. Temporally, the population was found to be maximum during J anuary, followed by J uly and minimum during Octo- ber. In general, the living specimens were found to be more in the earlier two stations, moderate in the in- termediate stations and lesser in the last four stations. Thus, one of the controlling factors of this species in the present area was depth. The optimum conditions for the abundance of living population of P. nip- ponica are: temperature 26.4 - 26.8C; salinity 30.2 - 30.8 %o and dissolved oxygen 2.8 - 3.6 ml/l. Osangularia venusta (Brady)The living speci- mens of O. venusta were more concentrated in the intermediate stations and were absent in station 13 and minimum in stations 1, 11, and 12 during any season. Seasonally, living population size showed a maximum during summer (April - 6271 specimens) followed by southwest monsoon (J uly - 5611 speci- mens). The active reproduction of the faunaseemed to occur irrespective of the season. A correlation of O. venusta population with different ecological pa- rameters measured shows that this species is tolerant of extreme changes of ecological conditions. The distribution and ecology of widespread and abundant living foraminifera revealed that among the eight species, Ammonia beccarii is found to have maximum reproduction during southwest monsoon and Pararotalia nipponica during winter. For Osangularia venusta, the active reproduction seems to occur irrespective of the season. The other five spe- cies reproduce actively during summer. In general, in the present area, the increase in temperature, salinity and dissolved oxygen content of the bottom waters are causative factors for the abundance of the living population.
Acknowledgement The authors thank the authorities of Fisheries De- partment, Tamil Nadu for providing motor launch to collect samples for the present work (Ph.D work of the first author). They also express their gratitude to the Head, Department of Geology, University of Ma- dras, Chennai and the Principal, National College, Tiruchirapalli for the permission to utilise the labora- tory facilities.
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