Indian Journal of Geo Marine Sciences Vol. 46 (04), April 2017, pp. 698-706
Experimental study on the regeneration of Fungiid corals in 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 Blair744 102, Andaman and Nicobar Islands, India *[E-mail:
[email protected]] Received 17 June 2014; revised 11 November 2016 Eleven species of fungiid corals were examined in situ condition for a period of 14 month after the mechanical excision to record their regenerative capability. Healing and structural development or regeneration was started within 1-6 months depending on the species and the pattern of excision of the coralla. Mushroom coral showed a tendency to regain its morphological symmetry through regeneration. Mouth took active part for the regeneration by organizing repair mechanism from the central direction while new mouth was formed in the absence of parental mouth for the survival of any part of coral fragments as well as with the existing one. [Key words: Regeneration, Fungiid corals, Andaman and Nicobar Islands]
Introduction Living coral tissues are the backbone to make competition1. Damages in coral reefs can be seen due to presence of variety of disturbances. Reefs can be destroyed in the form of rubble like structure by the active effect of Tropical storms2-5. There are a number of influencers such as predators, disease and low tidal effect which have the ability to remove the live tissues of hermatypic corals6-8. The decline of scleractinian lives reported worldwide over last two decades due to natural and anthropogenic factors2, 9-15. Among all, the corals under the family fungiidae are most common in shallow water regions throughout Indo-Pacific zone16. Free living stage in the lifecycle with the attached form can be seen in several individuals of this family. This mode helps them to continue their life stage on the different substrata like rubbles, sand and with other reefs also17-19. There are a number of causes viz. wave action, wind flow, sedimentation, temperature changes, emersion at low tide and freshwater flash flood can make damage on the shallow water corals20-24. The damages can be recovered by the two major processes such as colonization of planktonic coral larvae and the rapid tissue, skeletal repair and regeneration of remaining fragments25-28. The repair mechanism of corals is
an important feature for their survival as well as building resistance against diseases7, 26, 28, 29 .Regeneration is the process which includes development of calcification in corals skeletal structure with presence of environmental parameters30.Regeneration processes of corals help the coral communities for the successful establishment2, 4, 31.Success rate of regeneration activity is solely depend upon the corals natural resilience to repair the damaged tissues and skeleton structures 7, 28, 29.Innate ability to regenerate against damage is dependent on species specificity and the nature of damage30. Fungiids are most common to the every reef environment of Andaman and Nicobar Islands. The capacity of regeneration is one of the prime factors to make them available in the reef environments32.Morphological structures with relatively thin disk like shape and solitary mode of life are prone to fragmentation due to presence of wave action and other conflicts at the residing area16, 33. It is not obvious that, regular regeneration can be taken place in all the species, but it can be seen mostly to the polystomatous species. Fragmentation process is the indicator as a method of reproduction for the scleractinians34. Regeneration is the specific ability of corals especially Fungiids. After breakage of skeleton
INDIAN J. MAR. SCI., VOL. 46, NO. 04, APRIL 2017
portion, it can be grown as the extended part of the same species which may help to build or establish framework of coral reefs. This present paper dealt with the regeneration process of several species of fungiids after the mechanical excision. Materials and Methods The field experimental study was made at Pongibalu Jetty (Lat.11°30.956’N and Long.092°39.206’E) of South Andaman, Andaman and Nicobar Islands. Study was conducted in situ condition at the depth of 4m. Eleven species of mushroom corals such as Fungiapaumotensis Stuchbury, 1833; Fungiafungites (Linnaeus, 1758); Fungiarepanda Dana, 1846; Fungiaklunzingeri Doderlein, 1901; Fungia corona Doderlein, 1901; Fungiascruposa Klunzinger, 1879; Ctenactiscrassa (Dana, 1846); Polyphylliatalpina (Lamarck, 1801); Herpolithaweberi (Houttuyn, 1772); Herpolithalimax Horst, 1921 and Cycloseriscostulata (Ortmann, 1889) under the five genera were selected to make the experiments. Confirmation of species identification was completed depending on the photographs in conjunction with Veron and Pichon35, Hoeksema16, Veron36 and Venkataraman et al.37. All the above said live samples were collected from the study area by employing SCUBA diving and kept primarily in a net bag in the experimental site. Corals were excised by circular rock saw (BOSCH, GDC 34m, F 002 G30 110, 000017314) with the presence of continuous flow of seawater. The steady water flow will prevent the excessive generation of heat during the excision as well as it will avoid tissue drying. The excision was made in various patterns such as-half cut through the mouth in vertical and horizontal plane, two terminal parts for elongated coralla, only terminal part of coralla, cone shaped part from mouth to perimeter, peripheral part only etc. All the fragmented specimens were tagged with individual identity number to make proper study on them and kept in the experimental plot for in situ observation by SCUBA diving. All the experimental specimens were kept in natural habitat within hollow, cylindrical enclosure with a diameter of 1m. Observation and monitoring was made on regular interval for a total of 14 months from December 2011 to January 2013 to record the regeneration pattern of fungiid corals. Photographs were taken by a digital camera (Sony Cyber Shot, DSC-T900, 12.1 megapixels, marine pack). After the completion of study period, all the
699
live samples of fungiids were collected and kept in freshwater for 5- 7 days 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. Morphometric measurements of regenerated parts were made with the help of verniar calipers (Aerospace, 074 15376) and centimeter scale. Corallites of the specimen were studied in detail for morphological features. Results Eleven species of corals were studied for an experimental regeneration in their natural habitat. Solitary free livings as well as colonial fungiids were considered for conducting this experiment. Coralla were excised (Table 1) carefully in various manners to observe the regeneration pattern of them. Excision was made from single to multiple as described in the table 1. The cuts were made mainly in two patterns. One was made along the radiating septa of the fungiids and the other was made in perpendicular to the septa (Plate 1). The skeleton of the fungiids healed within a period of 10-20 days from the day of excision. As the individuals were kept in the natural habitat, exposed cut portion of the corals were partially covered with several fouling organisms like algae, tunicates, sponges etc. The aggregation of the fouling organism was noticed within 30 days of the experiment and persisted for a long time on many of the specimens. All the excised corals, used for experimental studies were survived throughout the study period of 14 months (Plate 2). The process of development can be described according the following categorizes. The septa of excised portion began to regenerate within 2 months if that portion has mouth and the other specimens started to regenerate mouth and axial furrow with that time. The mouth can be seen as the terminal part due to vertical and horizontal excision. The terminal portion of mouth started to develop septa for the completion of existing mouth as well as to form multiple mouth or extra mouth within the period of 4 months. The portion which had whole mouth did not form any extra mouth during the regenerative developmental process. Incised portion of corals without mouth started to form multiple mouth or extra mouth to make it as complete one within the period of 4 months. In some corals, it was seen that anthoblast was formed near the mouth portion and got detached from an anthocaulus. The development of axial furrow and corallites showed a well developmental stage for the
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MONDAL & RAGHUNATHAN: EXPERIMENTAL STUDY ON THE REGENERATION OF FUNGIID CORALS
colonial fungiids. Septa and the mouth showed a smooth development of their structural organization with the period of 180 days. The mouth showed its development as a complete mouth with the period of one year and complete new corallites (in case of colonial fungiids). Observation was made for next two months to find out their regeneration pattern after the complete development of mouth and axial furrow and corallites. Deviations in regeneration pattern are observed among the specimens and were examined morphometrically to make comparative account among the species studied. The process of regeneration was seen in the form of development of skeletal part of the scleractinians. Structure, shape, size are the features to control the regeneration pattern. Small portion of coral showed a higher degree of regeneration than the larger one (Table 1). Regeneration pattern of 11 species of corals were quantified in unit length development to compare inter species (Fig. 1). It was seen that the colonial species Polyphylliatalpina showed highest (0.333cm) development in unit length whereas the Fungiascruposa showed the lowest (0.033cm). All the species under the genus Fungia showed a less variable range of regeneration pattern. The values are very close to each other. Additions of skeletal part as the form of regeneration were clearly documented while the lighter colour tissue and the septal part were seen as non-parallel to the polyp. Intra species development of regeneration was Inter species regeneration of fungiids per unit length0.333 (cm) 0.234 0.226 0.219
0.191
0.236
0.12 0.033
coralla where only one terminal part was cut (without damaging the mouth) (Fig. 2). In Fungiafungites and Fungiarepanda maximum regeneration was seen in the portion of corals where cut was made through the mouth (1/2 mouth) (Fig. 3 and 4). Presence and absence of mouth play a crucial role for the regrowth of fungiids. The usual higher level of development was observed among those which had mouth by any means. New growth of septa in fan shaped structure was formed from the mouth region. Development in skeleton could be found from the inner septal edges as well as outer septal edges. Inner septal edges form mouth parts whereas outer septal edges form polyp margins. Intra species regeneration at per unit length (cm) of F. paumotensis
0.38 0.212
0.21
0.244 0.124
Through 1/2 2 Terminal Parts 1 Terminal Part Mouth
Through 1/4 Mouth
Terminal portion without mouth
Fig. 2- Intra species regeneration of F. paumotensis Intra species regeneration at per unit length (cm) of F. fungites
0.238 0.2
0.222
0.066 0.074
Terminal part
Through 1/2 Mouth
Fig. 3-Intra species regeneration of F. fungites Intra species regeneration at per unit length (cm)
0.2
Fig. 1- Inter species regeneration pattern of fungiids
also observed in the 3 species out of 11 depending upon the cut direction. With the extent of observation, it was seen that portion of coralla without mouth of Fungiapaumotensis showed higher degree of regeneration in skeleton length whereas the minimum was seen in the part of
0.183 Through 1/2 Mouth
Through 1/4 Mouth
Fig. 4- Intra species regeneration pattern of F. repanda
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INDIAN J. MAR. SCI., VOL. 46, NO. 04, APRIL 2017 Table 1- Experimental fungiid corals and their regeneration length Sl. No.
Specimens
Excised Position
Mouth/ Axial furrow Position in Coralla
Length during (cm) excision
Length after Regeneration (cm)
Regenerated Length (cm)
Mouth/ Axial furrow Status
1.
Fungiapaumotensis
1/2 Mouth
Terminal
4.9
6
1.1
Mouth completed
1/2 Mouth
Terminal
4.5
5.8
1.3
Mouth completed
1/2 Mouth
Terminal
5.6
6.4
0.8
Mouth completed
2 terminal parts
Central
5.7
6.9
1.2
No changes
1 terminal part
Beside terminal
6.9
7.8
0.9
No changes
1 terminal part
Beside terminal
7.6
8.8
1.2
No changes
1 terminal part
Beside terminal
8.6
9.4
0.8
New mouth
5
1/4 Mouth
Terminal
4.9
6.1
1.2
Mouth completed
1
Only terminal part
No mouth
3.1
4.3
1.2
New mouth
1/2 Mouth
Mouth terminal
4.1
5.2
1.1
Mouth completed
1/2 Mouth
Mouth terminal
6.9
8.2
1.3
Mouth completed
1/2 Mouth
Mouth terminal
3.6
4.5
0.9
Mouth completed
Terminal part
No mouth
7
8.4
1.4
New mouth
1/2 Mouth
Mouth terminal
6.7
8.3
1.6
Mouth completed
1/2 Mouth
Mouth terminal
6
7.2
1.2
Mouth completed
1/4 Mouth
Mouth terminal
5.3
6.3
1
Mouth completed
1/4 Mouth
Mouth terminal
5.6
6.6
1
Mouth completed
2.
3. 4.
Fungia corona
Fungiafungites Fungiarepanda
5.
Fungiaklunzingeri
1/2 Mouth
Mouth terminal
5.5
6.8
1.3
Mouth completed
6.
Fungiascruposa
1/2 Mouth
Mouth terminal
8.9
9.2
0.3
Incomplete
7.
Ctenactiscrassa
1/2 Axial furrow
Axial furrow terminal
11.9
13.2
1.3
Axial furrow not extended
1/2 Axial furrow
Axial furrow terminal
6.9
8
1.1
Axial furrow not extended
8. 9.
Herpolithaweberi Herpolithalimax
Central Near central
Terminal and Elongate Terminal and Elongate
10.6 22
11.3 22.6
0.7 0.6
Incomplete
Near central
Terminal and Elongate
14.1
15.4
1.3
Middle portion
Terminal and Elongate
6.2
6.8
0.6
Portion 1/2 Mouth
Entire portion Mouth terminal
3.6 3.6
4.8 4.4
1.2 0.8
10. 11.
Polyphylliatalpina Cycloseriscostulata
Axial furrow extended Mouth completed
Extra New Mouth formation
4
4
1
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MONDAL & RAGHUNATHAN: EXPERIMENTAL STUDY ON THE REGENERATION OF FUNGIID CORALS
Plate 1- Experimental excision of fungiids a-Fungia repanda (Excised through half of the polyp); b- Fungia paumotensis (Excised one terminal portion); cFungia paumotensis (Excised two terminal portions); d- Fungia repanda (Excised through half of the polyp); eFungia paumotensis(Excised through half of the polyp); f- Fungia paumotensis(Excised through half of the polyp); g- Fungia klunzingeri (Excised through half of the polyp); h- Fungia paumotensis (Excised one terminal portion); iFungia klunzingeri (Excised through half of the polyp); j- Herpolitha weberi (Excised through the central portion).
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Plate 2- Regeneration of fungiids after 420 days Complete development of excised mouth, extra mouth structure and of septal portion of different mushroom corals a- Fungia paumotensis; b- Fungia paumotensis; c- Fungia paumotensis; d- Ctenactis crassa; e- Fungia fungites; fFungia fungites; g- Polyphyllia talpina; h- Fungia paumotensis; i- Ctenactis crassa; j- Herpoltha weberi; kHerpoltha weberi; l- Herpoltha weberi
703
704 MONDAL & RAGHUNATHAN: EXPERIMENTAL STUDY ON THE REGENERATION OF FUNGIID CORALS Discussion Regeneration denotes the process of calcification to recover the fractured portion. Through the process live tissue can be seen as the regrowth form over the fractured or damaged area. Thus the recovery of live tissue can be seen with this procedure25, 27, 38. Volatile environmental conditions show the greater diversified fungiids to live with the sediments and other factors39. The environmental stresses helped them to acquire such a kind of property of regeneration to combat for their survival. Morphological attributes are one of the significant characters of fungiids to build the tissue-skeleton relationship to make the regeneration procedure effective in a progressive manner. It was found that the regeneration of fungiids were more rapid at the septal sides of the experimental organisms. The recovery of the corals is dependent on species specificity. Recovery is comparatively fast in branching corals25 where as low in massive corals38 as the exposure of skeleton matters. The corals were recovered and regenerated in a healthy state within the period of 30 days to 180 days. The development of the regenerated part became strong architecture with a period of one year or next six month of the mouth formation as well as recovery of wounds. Small portion of corals showed rapidity in growth in comparison with the large fragments of corals. Solitary corals have the ability to show its structural symmetry through the process of regeneration27. With the analysis of the present study, it was noticed that fungiids showed the tendency to regain its lost symmetry by the process of regeneration. Pattern of cut or breakage showed variation in skeletal regeneration. The study implies that the regeneration of solitary corals regenerate the broken part from a single polyp whereas the colonial fungiids from corallites. Skeletal development is usually prominent and undertake in the septal regions. Regeneration is along the inner septal edges in the mouth region. Structural development of regeneration pattern among the experimentally and naturally broken solitary fungiids is very closely related or appears to be similar27, 40. Study reveals that Ctenactiscrassado not show extension axial furrow from the fragmented area. Regeneration completes the structure of the species by developing the septal edges as a means of development whereas in colonial
Polyphylliatalpina, extension of axial furrow was recorded with presence of new corallites which were formed due course of regeneration procedure of that species. Mouth can be encountered as the controlling force of regeneration by organizing repair mechanism from the central direction. Development of new mouth can be formed in the absence of parental mouth for the survival of any part of coral fragments. In the present experiment, it was seen that all the excised portion of experimental samples showed regeneration by the development of mouth, extra mouth, septal or skeletal formation. The portions without parental mouth generate new mouth. Sectioning of samples is another important part of this experiment which controls the pattern of expansion. Cut portion without mouth, regenerate multiple mouth along the cut edges. With this experimental outcome it can be said that mouth is the most important part of fugiid corals for their development and play a key role to monitor the central organization role for the controlling of regeneration as well as normal developmental process. Due to specified morphological attribute such as solitary, freeliving (most of them), shape, size etc., the fungiids are used as the nuclei of the any reef areas. Formation of new reef areas can be initiated with these corals. The vulnerability in shape or structural pattern put them under high risk of breakage by means of natural as well as anthropogenic threats. The above said ability i.e. regeneration of fungiids safeguard the corals from the threat of breakage which leads to depletion of corals. And also it helps the corals for making survival to carry out its biological functions through regeneration. Conclusion The experimental studies on regeneration attributes of the fungiid corals were investigated in Andaman and Nicobar Islands. Eleven species of the mushroom corals were selected form the said studies. All the model animals exhibited regeneration through the process of healing and fouling. With the tendency of gaining morphological symmetry, the skeletal of the corals developed by the secretion of aragonite structure. Formation of mouth was recorded as the crucial developmental step during regeneration process and these formations controls the developmental process of
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regeneration for the mushroom corals. Presence of multiple mouth expedited the process in comparison with single mouth. Acknowledgements Authors are grateful to the Director, Zoological Survey of India, for providing necessary facilities to carry out the experiments. Authors are also thankful to the Ministry of Environment, Forest and Climate Change, Government of India for providing financial assistance to undertake the study through the projects of National Coral Reef Research Institute, Zoological Survey of India, Port Blair. The logistic support extended by Department of Environment and Forests, Andaman and Nicobar Administration is duly acknowledged. References 1.
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