Age and growth of fungiid corals of Andaman and Nicobar Islands, India

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1 Indian Journal of Geo Marine Sciences Vol. 46 (8), August 217, pp Age and growth of fungiid corals of Andaman and Nicobar Islands, India Tamal Mondal * & C. Raghunathan Zoological Survey of India, Andaman and Nicobar Regional Centre, National Coral Reef Research Institute, Haddo, Port Blair , Andaman and Nicobar Islands, India * [ t_genetics@yahoo.com] Received 25 February 215 ; revised 16 November 216 Two species of free-living and solitary corals such as Fungia paumotensis Stuchbury, 1833 and Fungia repanda Dana, 1846 under the family Fungiidae were examined inside in situ condition for 2 years to correlate and compare the age and growth relationship in Andaman Sea. Quarterly analysis of data revealed that, Fungia paumotensis showed higher growth rate {(.323 ±.28 cm (mean ± SD)} while Fungia repanda showed lower growth rate {.236 ±.45 cm (mean ± SD)}. Fungia paumotensis registered lower correlation co-efficient (R² =.422) with the age whilst Fungia repanda shown higher correlation co-efficient (R²=.857) with the age. The ratio of morphological attributes remained same during the course of aging for both the species. Mass of dry skeletal amplified exponentially with the length of the polyps for both the experimental species. Fungia paumotensis attained more skeletal mass than Fungia repanda during the size increment. Growth rings at the aboral sides of the coralla indicated a maximum age of 7 years i.e. post-tsunami recruitment of Andaman and Nicobar Islands. [Key words: Age and growth, fungiidae, scleractinian corals, Andaman and Nicobar Islands] Introduction Most of the scleractinian corals are restricted to tropical waters of the globe in its species content. Some of the scleractinians are adapted towards the temperate waters for their survival 1. Growth rate of scleractinian coral species is the main component for reef formation. Larger corals have higher reproductive success and also have superior competitive advantage 2. The pattern of coral growth is closely related to rate of calcification controlled by a number of factors such as light, temperature, feeding and the saturation state of the sea water. The calciumcarbonate saturation state is the ratio of the ion concentration product to the solubility product for the mineral deposited during a given time. Rate of chemical precipitation is proportional to calciumcarbonate saturation. Greater the concentration of ions, the greater is the formation of mineral 3. The rate of growth pattern and sometimes depends on the growth of zooxanthallae by the animal host via nutrient limitation, which is known as a paradox 4-6. Process of calcification in hard corals is a biogenic action firmly maintained via an organic matrix 7, it is strongly affected by the energetic status of the coral. The activity is regulated both by the photosynthetic activity of the endo-symbionts and by the feeding command of the host 8, 9. The presence of light and type of food availability has a species-dependent consequence on rate of calcification. Life of scleractinian corals starts from the settlement of planula larvae. Larval survival, recruitment of juveniles, growth of somatic, reproduction through sexual process and the mortality of scleractinians is variable in wide state from species to species which monitor the development of reefs through the way of succession 1, 11. Some scleractinian corals follow r-strategy as they produce large number of propagules during the reproductive season of each year. The growths of those are very rapid and take pioneer role to settle recently formed space on reef 1, Other scleractinians follow k- strategy as they reproduce during a short period each year and the growth rate is slow. Though the corals take several years to make settlement of fresh substrata, the successful establishments are able to dominate the reef through the effective defense of living space 1, 15, 16. Growth patterns of the corals are

2 INDIAN J. MAR. SCI., VOL. 46, NO. 8, AUGUST variable according to species and that hardly ever experience fission or fuse while partial mortality can be noticed by distortions of the usual growth appearance among stony corals with compact branching colonies 12, 17, 18, massive hemispherical 15 and free-living fungiids 19 etc. If process of fractional mortality and fragmentation of the corals do not take any part as interference, normal growth can take place indeterminately. 11, 15, 17, Age is a linear purpose of individual length 18. According to von Bertalanffy (1938) 19, the growth pattern of reef building corals are actively associated or related with the age of the corals. Age and size have a close relationship. Changes in stony corals and pattern of turnover are explicable with the growth model which can be expressed by age-size relationship. Fungiidae is the most important family of scleractinian corals due to presence of solitary appearance among the others with the unique form of free-living coral polyps. These corals are susceptible to overexploitation. With the presence of mobile activity, the only free- living fungiids move off the reef and settle down in sandy areas of reef associated environment. These settlements of corals in new areas serve as the nuclei for the formation of new reef patches 2. Formation of any reef area can be initiated with the presence of mushroom corals at reef flats and slopes 21, 22. Asexual reproduction helped the coral for rapid recovery after disturbance 23 though sexual reproduction is also a significant event for the survival 24. The asexual activity accelerates the formation of large patch reefs during a short period with only the fungiidae corals 25. Present paper dealt with the analytical description of growth pattern on two species of fungiidae corals of Andaman and Nicobar Islands. Material and Methods The study was carried out at Pongibalu Jetty (Lat N and Long E) of South Andaman at the depth of 7 m. Fungia paumotensis Stuchbury, 1833 and Fungia repanda Dana, 1846, the two species under the same genus were selected for the study. Undersea growth studies and regular monitoring were made by SCUBA diving. Undersea morphological measurements on growth were made with the help of Vernier calipers (Aerospace, ) and centimeter scale. Growth data were collected at 3 months interval following the procedure of Chadwick-Furman et al. 26. Data were collected on the length of fungiids along their mouth axis and width which is perpendicular to the mouth axis 27, 28. All the experimental specimens were kept in a net made enclosed area of reef slope to restrict their movement to control missing data. Observation was made for a period of 24 months to document the growth rate of the fungiids. After the completion of study period 3 live samples of Fungia paumotensis and Fungia repanda were collected and kept in freshwater for one week with 2-3 changes of water. After that the specimens were thoroughly washed in freshwater to get the calcareous skeleton structure of calcium carbonate and kept for sun drying for a period of 1 days. Dry mass of all the samples were measured by using pan balance (Docbel, BARUN). The ring structure at the ventral side of the fungiidae corals were studies as those correspond to the annual growth 26, 27. Length frequency method was applied to demonstrate the multimodal length distribution of a population of different age groups 29. Growth rate of the fungiidae corals were also determined by von Bertalanffy s growth curve and Ford-Walford method, computerized non-linear regression method using SPSS software 19. Results 3 live specimens of Fungia paumotensis and Fungia repanda were selected to get quantitative data of their growth rate. There was no report of mortality of the experimental animals during the study period. The data of growth rates were analyzed at 3 months interval for the period of 2 years. Fungia paumotensis Stutchbury, 1833: Quarterly mean growth rate of Fungia paumotensis was.323 ±.28 cm. The width of the corals varied linearly with the length of the corals (Fig. 1). Dry mass of the individual coral samples was analyzed in accordance with the length of those. Mass of the samples increased exponentially with the length (Fig. 2). A total of 12 coral polyps were measured to study the morphometric growth pattern of the fungiids. Morphology of the 12 coral polyps were elongated or slightly oblong in shape which can be justified with the ratio of width and length (.77 ±.167). Relationship of width and length in morphological character i.e. width and length ratio of the polyps were constant (Fig. 3). The constancy of the ratio indicates the regularity of their structure plane without any changes in future of their life span. This is the isometric growth pattern of life. The size of the corals increased with the time. It depends on the initial stage. Growth rate of this species decreased linearly with the increased coral size (Fig. 4). As the

3 Coral width (mm) Width: Length Dry skeletal mass (g) 1634 MONDAL & RAGHUNATHAN: AGE AND GROWTH OF FUNGIID CORALS documentation was recorded quarterly, it was observed that maximum growth (.37 cm) was noticed during the 3 months periods of June to August 211 whereas minimum (.28 cm) was recorded during March to May 212. Physical observation of the coral polyps indicated the 7 years maximum age of the corals depending on their ventral growth rings (Fig. 5). Fungia repanda Dana, 1846 Quarterly mean growth rate of Fungia repanda was.236 ±.45 cm. The polyps of this species are circular shaped mostly. Depending on the shape of this species, only diameter was documented to observe and analyze the growth pattern. Dry mass of all the 3 individual coral samples were recorded and analyzed in comparison with the diameter of those. It was observed that the mass of this species increased exponentially with the diameter (Fig. 6). As two species were considered to carry out experiments, diameter was described as length to make comparative analysis during inter-species relationship y = 6.497x R² = Coral length 1 (mm) 15 2 Fig. 2. Exponential relationship of coral length and dry skeletal mass of Fungia paumotensis N=12 y =.724x -.86 R² = N= Coral length (mm) Fig. 1. Coral length and width linear relationship of Fungia paumotensis Coral length (mm) Fig. 3. Graph showing polyp width: length ratio of Fungia paumotensis

4 Dry skeletan mass (g) Growth rate (cm/3months) INDIAN J. MAR. SCI., VOL. 46, NO. 8, AUGUST y = -.7x R² = Observation for 2 years in each 3 months interval Fig. 4. Size dependent coral growth rate of Fungia paumotensis Coral showed a same morphological structure i.e. circular shaped during the entire lifespan. The size of the corals increased depending on the age. Growth rate of this species decreased linearly with the increased coral size (Fig. 7). It was also observed that maximum growth (.3 cm) was seen during the 3 months periods of June to August 211 whereas minimum (.19 cm) was seen during the two periods such as September to November 212 and December 212 to February 213. Growth rings were observed on the ventral sides of the polyps. It was seen that the maximum age of this species was 6 years (Fig. 8). Non-parametric methods ELEFAN Optimized growth parameters (L and K) and the goodness of fit index (Rn) was obtained for growth parameters of Fungia paumotensis and Fungia repanda by ELEFAN I method in the FISAT II package. The non-seasonalized length frequency histograms with growth curves for F. paumotensis and F. repanda are shown in Figs. 9 and 1. Automatic search routine in FISAT package derived the L and K values of mm and 1.5 yr -1 for F. paumotensis and and.51 yr -1 for F. repanda respectively, whereas the K-Scan routines gave the values of mm and.34 yr -1 for F. paumotensis and mm and.69 yr -1 for F. repanda respectively y = 3.169x R² = Fig. 5. Aboral view of growth rings of coral polyps as indicator of annual growth (a. Fungia paumotensis- age 6years; b. Fungia paumotensis- age 7 years; Fungia paumotensis- age 7 years; Fungia paumotensis- age 7 years). Each ring corresponds to a year of growth. Scale is in cm. Coral length (mm) Fig. 6. Exponential relationship of coral length and dry skeletal mass of Fungia repanda

5 Growth rate (cm/ 3months) 1636 MONDAL & RAGHUNATHAN: AGE AND GROWTH OF FUNGIID CORALS.35.3 y = -.17x R² = Fig. 9. ELEFAN I growth curve of Fungia paumotensis Observations for 2 years in each 3 month interval Fig. 7. Size dependent coral growth rate of Fungia repanda Fig. 1. ELEFAN I growth curve of Fungia repanda Estimation of growth parameters from length at age data - The results of fitting growth parameters L, K and t o to von Bertalanffy growth equation for obtaining length at age are given in Table 1 for F. paumotensis and Table 2 for F. repanda. Table 1. Fitting of von Bertalanffy growth equation for Fungia paumotensis L = mm K =.34 t o = t 1-e -K(tto) (yr) t-to k(t-to) e -K(t-to) Lt = L [1- e -K(t-to) ] Fig. 8. Aboral view of growth rings of coral polyps as indicator of annual growth (a. Fungia repanda- age 6years; b. Fungia repanda - age 6 years; Fungia repanda - age 5 years; Fungia repanda - age 6 years). Each ring corresponds to a year of growth. Scale is in cm

6 Lt+1 Lt + 1 INDIAN J. MAR. SCI., VOL. 46, NO. 8, AUGUST Table 2. Fitting of von Bertalanffy growth equation for Fungia repanda L = mm K =.69 t o = Y=.513x t (yr) t-to k(t-to) e -K(t-to) 1-e -K(tto) Lt = L [1- e -K(t-to) ] Lt Ford-Walford method Graphical depiction of Lt+1 on Lt is shown in Fig. 11 for F. paumotensis and Fig. 12 for F. repanda to estimate the asymptotic length. The regression line fitted to the points showed the following equation for F. paumotensis Y=.7121x and Y=.513x for F. repanda. The functional regression slope, b, and intercept, a were.7121and 4.69 in the case of F. paumotensis and.513and 9.89 in the case of F. repanda respectively. As the slope of the line b is related to the growth coefficient K, the value of K was deduced using the following equation: K = -ln [b ] The value of L was deduced by employing the following equation: L = a / (1-b ) The estimated K values for F. paumotensis and F. repanda by Ford Walford method were.34 yr -1 and.69 yr -1 for F. paumotensis and F. repanda respectively. The estimated L values were mm and respectively Y=.7121x Fig. 12. Ford-Walford plot for Fungia repanda von Bertalanffy plot The results of von Bertalanffy plot for the estimation of K and t o for males and females of F. paumotensis and F. repanda are shown in Tables 3 and 4 and are graphically depicted in Figs. 13 and 14. The L values estimated by the Ford Walford method (16.28 mm for F. paumotensis and mm for F. repanda) were used as input. The values of the regression coefficient a, obtained were.213 for F. paumotensis and.248 for F. repanda. Slope b, which is equal to the K value, was.34 yr -1 for F. paumotensis and.39 yr -1 for F. repanda. The t o values estimated for F. paumotensis and F. repanda were -.489yr and -.36yr respectively. Table 3. Estimation of K and t with the von Bertalanffy plot for Fungia paumotensis Age Length (Lt) Lt/L 1-Lt/ L -Ln(1- Lt/ L ) K=b=.34, t = (-a/b) = Table 4. Estimation of K and t with the von Bertalanffy plot for Fungia repanda Lt Age Length (Lt) Lt/L 1-Lt/ L -Ln(1- Lt/ L ) k=b=.69, t = (-a/b) = -.36 Fig. 11. Ford-Walford plot for Fungia paumotensis

7 -Ln(1-Lt/Lα) -Ln(1-Lt/Lα) 1638 MONDAL & RAGHUNATHAN: AGE AND GROWTH OF FUNGIID CORALS Age Fig. 13. von Bertalanffy plot for Fungia paumotensis Y=.69x Fig. 14. von Bertalanffy plot for Fungia repanda Y=.34x Age Discussion The selected animals for the study were freeliving during adult stage, solitary in nature. There was no report of mortality during the study periods indicate the experimental site is environmentally stable. As most of the reefs building corals are colonial, growth can be seen as long unlimited process depending on the body size 11, 3. Previous experimental data shows that several hard corals are known for reducing their growth rate parallel to their age. Among the freeliving fungiids several corals such as Ctenactis echinata, Diaseris distorta, Fungia actiniformis, Fungia fungites and Fungia granulosa showed age dependent the growth rate which reduced gradually 26, 28, Present study supports the previous work after Bablet 1985 regarding age and growth rate relationship and also depicts that the growth pattern of Fungia paumotensis and Fungia repanda is age dependent and reduced gradually as they aged. Though the growth rates of both the species are dependent on same variables, but Fungia paumotensis shows comparatively higher quarterly growth {(.323 ±.28 cm (mean ± SD)} with lower correlation co-efficient (R² =.422) with the age and Fungia repanda shows lower quarterly growth {.236 ±.45 cm (mean ± SD)} with higher correlation co-efficient (R² =.857) with the age. The Fungia paumotensis is elongated in shape (width: length=.77). The species shows a gradual continuation in the same ratio. Polyps of the Fungia repanda are circular and show the same structural plane according to age. Dry skeletal mass of both the fungiids increased exponentially with the length of the polyps. Fungia paumotensis gains more skeletal mass during the size increment than Fungia repanda. Thus Fungia repanda shows relatively flat (R² =.863) exponential length-dry skeletal mass growth curve than the stiff (R² =.878) curve of Fungia paumotensis. In the present study a total of 12 polyps of Fungia paumotensis and 3 polyps of Fungia repanda, it was seen that the maximum age was 7 and 6 years respectively depending on the annual growth rings on their aboral surface. Growth rings are used as unique feature for the non-destructive aging of corals 26, 27. The extensive analysis of growth pattern depending on the growth rings, it can be said that the growth of the corals continued from the centre to the perimeter for both of the species. Though these are solitary free living corals, but the recruitment of the juveniles occur through the natural process of larval settlement. In 24, Tsunami resulted the destructive mode of life in every aspect of the coral through the alteration of sea water environment as well as by direct tremendous powerful wave action. Corals of all the families were damaged due the natural calamity. As of Fungia paumotensis and Fungia repanda are freeliving and solitary corals under the family fungiidae, it can be predicted that wave action of Tsunami destroyed the low weight fungiids (Fungia paumotensis and Fungia repanda) most. Settlement of juvenile corals needs solid, concrete substratum and dead reef 35-37, lower siltation rates, oligotrophic condition or lower algal biomass 38, lower light intensities 39, favourable temperature and salinity. The physical environment of the sea came in biogenic state after this massive alteration during the periods of next one year tentatively 4. Recruitment of the experimental species was

8 INDIAN J. MAR. SCI., VOL. 46, NO. 8, AUGUST occurred after the period of environmental settlement as the post tsunami recruitments. The maximum age of 7 years support the statement that the all the experimental animals were recruited after the tsunami in study areas of Andaman and Nicobar Islands. The present paper implies the growth pattern of the two fungiids as an initial frame work to understand the parts of development process of their life cycle. The study will be helpful to understand the life pattern of fungiids for their sustainable development and conservation of scleractinian corals in their own niche. Conclusion The experimental studies on two species of free living fungiid corals were carried to document their age and growth pattern in Andaman and Nicobar Islands at the depth of 7 m. Fungia paumotensis exhibited higher degree growth in comparison with Fungia repanda. The growth if the corals were proportion with the structural attributes of the specimens. 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