CEYLON J.SCI.(BIO.SCI.) VOL.17&18,1984-85 SOME ECOLOGICAL ASPECTS OF THE MANGROVES ON THE WEST COAST OF SRI LANKA * ** K.H.G.M. De Silva and S, Balasubramaniam ?Department of Zoology, University of Peradeniya, Sri Lanka. "Department of Botany, University of Peradeniya, Sri Lanka. ABSTRACT Mangrove formations (mangals) on the west coast are not extensive and, in most places, are limited to narrow strips bordering river mouths and estuaries. A clear ecological zonation is difficult to discern, except for a Rhizophora-Bruquiera zone at the edge of the water followed by a zone containing the other core species. The low level of tidal amplitude as well as persistent and long-term human interference are major factors that led to the limitation of mangroves to narrow strips and to the disappearance or non-establishment of proper ecological zonation. Floristic composition of wet zone mangroves is somewhat different from that of dry zone mangroves. This difference appears to be due to prevailing salinity differences, wet zone mangroves experiencing lower salinity conditions and dry zone mangroves experiencing higher salinity conditions. In each zone, there are riverine and estuarine mangroves. Four types of high salinity (dry zone) estuarine mangroves, viz. fringing, scrub, overwash, and basin, could be. distinguished depending on the geomorphic and flooding characteristics. Basin mangroves are not found on the west coast. Mangroves in Sri Lanka have been subjected to man's ' detrimental interference to a greater or lesser degree for a long time. In the north-east, mangroves have received 23 K.H.G.M. DE SILVA AND S. BALASUBRAMANIAM relatively less interference, but, those on the west and south? west coasts have been highly interfered with, and in some cases, mangroves have virtually disappeared. Urgent conservation measures are required to save Sri Lankan mangroves. INTRODUCTION Mangrove formations (or mangals, as some authors prefer to call these communities, reserving the term mangrove to individual plant species (Macnae, 1968)), are a prominent feature of the tropical coastal ecosystems and they develop in sheltered shores of lagoons and estuaries. They contain a characteristic assemblage of salt-tolerant trees and provide habitats for a variety of fauna and nursery grounds for juvenile finfish and shellfish. They produce large amounts of organic matter, which are used in food webs of lagoons and estuaries. They also help in the prevention of shore erosion. As Tansley & Fritsh (1905) pointed out at the beginning of the century, mangroves in Sri Lanka are confined to the muddy shores of the estuaries and are not found at all on the actual sea shores (which are sandy) in contrast to the extensive mangroves found in the shores of the Straits of Malacca? on the west coast of the Malay peninsula. Mangroves are perhaps the most misused coastal ecosystem in Sri Lanka, and through ignorance, people have destroyed most of this valuable natural resource. Even at present, for most of the general public, mangroves are an evil- smelling wasteland infested with mosquitoes and since they have only a very low "real-estate" value they should be reclaimed if possible and put to "better" use such as for growing crops and establishing human settlements. Mangroves in Sri Lanka received little attention even from ecologists until the last two decades. The present extent of mangroves in Sri Lanka has variously been estimated at over 4,000 ha (Arulchelvam, 1968) to over 6,000 ha (Jayawardene, 1985). Recently preliminary studies of mangrove flora in the Puttalam-Kalpitiya region (Amarasinghe & Perera, 1984), Vadamarachchi estuary in the north (Balasubramaniam, Rasiah & Ratnapala, 1970) and some of the islands in the mouth of Negombo estuary (Pinto, 1982) have been reported. Although virtually all macrophytes and most of the macrofauna of the Sri Lankan mangroves are known from various works, mainly of a taxonomic nature, little is known of the MANGROVES OF SRI LANKA l i f e cyc le s of most of the s p e c i e s . Studies on ecology, zonation and d i s t r ibut ion of mangrove spec ies in Sri Lanka are lacking except for those that are mentioned e a r l i e r . The present study deals mainly with the types of mangroves on the west coast of Sri Lanka, the i r eco log i ca l zonation and the e f f e c t of man's interference on them. MATERIALS AMD METHODS Mangroves in the r iver mouths of Gin Ganga, Kala Oya and Pomparippu Aru, and those on the shores of larger i s lands in Kalpi t iya , Negombo and Balapit iya e s tuar i e s as wel l as those on the mainland bordering these e s tuar i e s and Hikka- duwa estuary (F igs . 1, 2 and 3) were studied during 1985/86. Herbaceous and woody mangrove macrophytes in each mangrove were i d e n t i f i e d and t h e i r r e l a t i v e abundance est imated. Macrophytes in several transects in each mangrove were studied and t h e i r zonation noted. Samples of macrofauna such as crabs, gastropods, b iva lve s , hermit crabs, shrimps and prawns and f i sh were c o l l e c t e d from the mangroves as wel l as from the adjacent e s tuar ie s and r iver mouths, and were taken t o the laboratories of the University of Peradeniya for further s t u d i e s . Zooplankton were c o l l e c t e d from Kalpi t iya , Negombo, Balapit iya and Hikkaduwa e s tuar i e s by drawing a zooplankton net (mesh s i z e 335^um) through water from a slow running boat . In addi t ion , other mangroves on the west coast (Fig . 1) were a l s o examined and the macrophytes present in them were noted. RESULTS Mangroves on the west coast are not extens ive and are often l imited to narrow s t r i p s . The larges t mangrove area was seen in the Kala Oya-Pomparippu Aru e s tuar i e s in the north-west (Fig . 2 ) . I t s extent has been estimated as about 1,800 ha (Kanakaratne, Perera & Fernando, 1984) . On the other hand, the extent of the mangrove in Balapit iya estuary (Fig . 3) in the south west i s l e s s than 100 ha and that in the estuary of the Gin Ganga (Fig. 1) in the south i s even l e s s . The t a l l e s t t r ee s are seen in the Kala Oya mangrove with Rhizophora spp . , Bruguiera s:pp. and Avicennia spp. growing to 10-15 m. In a l l other mangroves, t r e e s growing over 10 m are except ional . K.H.G.M. DE SILVA AND S. BALASUBRAMANIAM 25 Although many tree species were commonly found in all mangroves examined, the floristic composition in the dry zone mangroves (e.g. those in Kalpitiya area) is somewhat different from that of the wet zone mangroves (e.g. those in Balapitiya area) (Table 1 ) . Nypa fruiticans is found only in the wet zone mangroves. Sonneratia caseolaris, which is found in the wet zone, is replaced by S_j_ alba in the dry zone. Rhizophora apiculata and Bruguiera sexanqula are common in the wet zone whereas R^ mucronata and B. gymnorhiza are common in' the dry zone. Dry zone mangroves often have saltmarsh plants such as Salicornia brachiata and Sueda spp. in the back mangrove whereas wet zone mangroves have mangrove associates such as Polichandrone spathacea and Annona glabra. Well developed beds of sea grasses such as Enhalus acoroides, Halophila ovalis, and Thalassia hemprichii are usually found in the fore mangroves in the dry zone. Mangroves in the Negombo estuary- are intermediate in character between the wet and dry zone types as expected since the Negombo lagoon lies in the border region between the two zones. In both zones, mangroves along the banks of river mouths were more luxuriant than those found in the other estuaries as the amounts of sediment deposited in the river estuaries are richer in nutrients than those deposited on the shores of the other estuaries. In order to distinguish between the two types, the mangroves on the river banks (or river estuaries) will be referred to as riverine mangroves and those on the shores of the other estuaries (including those on the shores of the islands in these estuaries) will be referred to as estuarine mangroves in this study (see also Discussion). The low tidal amplitude and the slow rise and fall of tides could not flush riverine and estuarine mangroves to any significant extent. However, during rains, riverine mangroves are flooded and flushed to a great extent with freshwater, followed by the deposition of silt and other materials brought by the rivers. Degraded scrub mangroves are occasionally seen in the dry zone (e.g. on shores of Mundel estuary) (Fig. 1 ) . Rhizophora" spp., Bruguiera spp. and Sonneratia spp., which are prominent in other mangroves, were absent in the scrub mangrove. Even dominant species such as Avicennia marina is often found to be stunted. These areas show poor drainage and get water-logged when they are occasionally covered by extreme high tides and during rains. Thus, the salinity of the soil varies considerably, and hypersaline conditions develop with evaporation. Overwash mangroves, which are flushed at each tide, were observed on small islands and small, low-lying finger-like processes of mainland projecting into the lagoon, usually perpendicular to the tidal flow. The floristic composition of the overwash mangrove is similar to that of the marginal zone of estuarine mangrove, but when the area of the overwash mangrove is very small, usually only R^ mucronata grew on it. Such overwash 26 MANGROVES OF SRI LANKA mangroves were seen at Keerimundel and Kovilkudah along the Kalpitiya estuary (Fig. 2 ) . A clear ecological zonation of different tree species was not observed except for a zone of Rhizophora spp. ( Bruquiera spp. and sometimes Sonneratia spp., at the edge of the water followed by a mixed zone containing other core species such as Avicennia marina, Aeqiceras corniculatum, Ceriops tagal, Excoecaria aqallocha and Lumnitzera racemosa. Bruguiera spp., are usually found behind Rhizophora spp-. Avicennia marina is sometimes found towards the upper edge of the mixed zone, and sometimes extending right onto the water's edge. Occasionally, small areas free of trees but covered with breathing roots of A^ marina were seen in t h e mixed zone. Lqmn.itzera 1 ittorea was recorded only from onfe locality in one of the small islands in the Balapitiya; estuary at the edge of the water. A list bf mangrove core species as well as associates of Sri Lanka is given in Table 2 . On the whole, there was a paucity of mangrove fauna, both in numbers and in diversity. The commonest were cerifhidean gastropods and grapsid crabs. Portunid crab, Scylla serrata was found in all estuaries examined. Other decapods included the ocypodid crabs Uca lactea and Macrophthalmus depressus, the anomurans Eupagurus sp., Pagurus sp. and Thalassina anomala, and several species of Penaeus and Metapenaeus. The bivalves Anadara spp., Geloina spp.., Gaf f rarium tumidum, Perna spp.. Pinna bicolor, and the gastropod Pleuroploca trapezium were found in the mud and in the beds of sea grasses. The oyster Crassostrea cucullata was seen attached to the roots and stems of; Rhizophora spp., as well as in mud. The common fish species in the estuaries included mugilids, cichlids, ambassids, latids and siganids. Periophthalmus koelreuteri was seen in the mudflats and on the prop roots of Rhizophora spp. Fish fry and fingerlings and juvenile prawns were often seen within mangroves in small, more or less permanent, water holes and small channels. Larvae of prawns and crabs, copepods and their nauplii were common in the zooplankton of all estuaries. However, some differences in the zooplankton composition of the wet zone estuaries and dry zone estuaries were observed, with freshwater species occurring in the estuarxes? in the wet zone and marine species occurring in those of the dry zone. K.H.G.M. OE SILVA AND S . BALASUBRAHANIAM 27 All mangroves were subjected to degradation by man to a greater or lesser degree. Very little mangrove was left in some of the estuaries such as at Hikkaduwa. In such areas, human settlements or crop plantations such as coconut, cinnamon and rubber extended almost to the edge of the water. Mangroves in many areas have been cleared for crop plantations from the early days of European rule of the coastal areas. For instance, cinnamon plantations were started after the Portugese landed in Sri Lanka in 1505. Such plantations could be seen in the areas of Balapitiya and Hikkaduwa estuaries (Fig. 3 ) . Even within the last 20 years, the mangrove cover of Negombo estuary (Fig. 2) dwindled very much (by more than 10% according to some estimates). Nypa fruticans which flourished in Hikkaduwa estuary about 50 years back (according to some eye witnesses) has now completely disappeared from there. Even during the period of the present study in 1985/86 some of the mudflats were prepared for cinnamon plantations. Among the many uses of mangrove trees in Sri Lanka, such as for firewood, timber, tanning purposes, fodder, use of branches of trees such as Rhizophora spp., Bruguiera spp. and Avicennia spp. for brushpile fishing in the Negombo estuary merits special mention. As branches are required every two or three weeks to make new piles, mangrove trees are regularly cropped for this purpose. However, this necessity has encouraged some people to replant, degraded areas with mangrove trees, although to a very limited extent. DISCUSSION Development of mangroves depends on the interplay of many factors, the most important of which are climate, geomorphology of the shore area, tidal amplitude, salinity and sedimentation characteristics. Mangroves develop in intertidal mudflats or humid sands in places where the substratum is not much disturbed by wave action (Abeywickrama, 1964). Geomorphology and tidal amplitude are of major importance in determining the extent and width of the mangrove, while the floristic composition of the mangrove in a particular area will depend, among other factors on the salinity prevailing in the region. In Sri Lanka, most of the coastal brackishwater bodies are called lagoons. However, according to the generaly accepted definition of Pritchard (1967) an estuary "is a 28 MANGROVES OF SRI LANKA semi-enclosed coastal body of water, which has a free connection with the open sea, and within which sea water is measurably diluted with freshwater derived from land drainage". Thus, an estuary is subjected to the influence of regular tidal fluctuations. On the other hand, a lagoon does not normally have a free connection with the open sea and may be inundated with sea water at irregular intervals only. Furthermore, a lagoon receives freshwater mainly- from direct rainfall rather than from land drainage (e.g. lagoon within an atoll). River mouths, coastal bays, tidal marshes, and bodies of water behind barrier beaches are cited by-Odum (1971) as examples of estuaries. .Thus, the majority, if not all, of coastal brackishwater bodies in Sri Lanka including the so-called lagoons (e.g. Jaffna lagoon, Kalpitiya lagoon, Negombo lagoon) would come under the category of estuaries according to Pritchard's definition, and in the present study they are considered as such. The tidal amplitude in Sri Lanka is small and rarely exceeds 1.0 m. This may have played a major role in restricting mangroves to narrow strips. However, Tennent (1859), more than a century ago, noted dense growth of mangroves on the shores of estuaries. This, by no means, is true today. The possibility of mangroves spreading to areas which are covered by tides very occasionally or not at all is evidenced by the luxuriant growth of some trees of Bruquiera qymnorhiza growing to about 10 m and Ceriops taqal growing to about 7 m, about 100 m away and about 2 m higher than the high water mark of the Negombo estuary. These trees were planted by man. It is also known that most mangrove species can be grown in freshwater (Mepham & Mepham, 1985). Thus, other factors would also have contributed significantly to restricting the mangroves in the west coast to narrow strips. Among these factors, human interference appears to be the most important which also would have caused the disappearance of the mangroves in some localities and the reduction of the extent of mangroves in the other localities to the present very low level. The relatively less interference seen in the Kala Oya mangrove can be attributed to the fact that it forms the south-west boundary of the Wilpattu National Park and is therefore strictly p r o t e c t e d . Five kinds of mangroves, namely, riverine, fringing, scrub, overwash and basin, are described in Sri Lanka depending on the topography, flooding characteristics and floristic composition. Balasubramaniam (1985) describes Nypa fruticans, Sonneratia caseolaris, Bruguiera sexanqula, Rhizophora apiculata and Dolichandrone spathacea communities as K.H.G.M. DE SZLVA AND S. BALASUBRAMANIAM 29 characteristic of the riverine type, and R.mucronata, B. qvmnorhiza, Avicennia officinalis, Aeqiceras corniculatum, Lmnitzera racemosa and E?coecaria agallocha communities as characteristic of the fringing type, and shows that riverine mangroves are present in river estuaries along the south and south west coasts, and fringing mangroves in Batticaloa, Rekawa and along other shallow estuaries. He also describes the floristic composition of the other types. Amarasinghe ? Perera (1984) recognize four types of mangroves, namely, riverine, fringing, scrub and overwash, in the Puttalam- Kalpitiya region (dry zone) on the topography and flooding characteristics. They consider the mangroves along the shore line of the mainland and of larger islands in the region as the fringing type, and those on the estuaries of KalaOya and Mi Oya as the riverine type. In the present study, no clear cut difference was observed in the floristic composition of the mangroves at river mouths and those bordering the other estuaries either in the dry zone or in the wet zone (Table 1 ) . However, the trees in the mangroves in the river mouths showed more luxuriant growth than those in the mangroves of other estuaries, as has been noted by others in other countries as well. It appears that the communities cited as characteristic of the riverine type by Balasubramaniam (1985) are characteristic of . the wet zone mangroves, and those cited as characteristic of the fringing type are characteristic of the dry zone mangroves. The floristic compositions of the two types appear to depend on the salinity prevailing in the estuaries. For instance, Nypa fruiticans, found in the wet zone, has a low salinity tolerance whereas Avicennia marina, found predominantly in the dry zone, has a high salinity tolerance. Moreover, the presence of the freshwater atyid shrimps (Caridina spp.) as well as the freshwater zooplankton species in the wet zone estuaries indicate the low salinity conditions prevailing in them. The presence of marine zooplankton species such as Caprella sp. in Kalpitiya estuary indicates the high salinity conditions prevailing there. High rainfall and high and fairly evenly distributed drainage from the surrounding land as well as from the rivers throughout the year make the wet zone estuaries less saline. The wet zone receives rains throughout the year but specially during the periods April-May and October-December, and the annual rainfall in the coastal wet zone ranges from 1,900 to 2,500 mm. The dry zone, on the other hand, receives rains mainly during the period October-December, and the annual rainfall in the coastal area ranges from 900 to 1,800 mm. The rains in the dry zone are very seasonal, whereas those in the wet zone are more evenly distributed. For each km 2 of catchment area the annual discharge ft.om rivers in the wet zone ranges from 4.8 x 10 to 2.4 x 10 m 3, whereas that from rivers in the 30 MANGROVES OF SRI LANKA dry zone ranges from 6.0 x 10 to 4 .8 x 10 m3 (Abeywickrama, 1964). Thus the mean discharge of the r iver Kalu in the wet zone would be 125 cumecs whereas that of Kala Oya in the dry zone (which has a catchement area s imilar in extent t o that of r iver Kalu) would be only 26 cumecs. I t i s perhaps bet ter to d i s t ingu i sh between low- s a l i n i t y (wet zone) and high s a l i n i t y (dry zone) mangroves on the f l o r i s t i c d i f ferences (as discussed e a r l i e r ) . Each of these types could be further divided into r iver ine and estuarine types depending on whether they border r iver mouths or other e s t u a r i e s . Four types of high s a l i n i t y - e s t u a r i n e mangroves could then be dist inguished depending on the geomorphic and flooding c h a r a c t e r i s t i c s . These are ( i ) the fr inging mangrove, which i s commonly found on the shores of the mainland bordering the e s tuar ie s and on. the shores of the i s lands in the e s tuar i e s ; ( i i ) the scrub mangrove, which appears t o develop under hypersaline condi t ions; ( i i i ) the overwash mangrove, which are flushed at each t i d e ; and ( iv ) the basin mangrove (which i s not present in the wet zone) which develops in low-lying b a s i n - l i k e e s tuar ie s ( e . g . Vadamarachchi estuary in the north (Fig . 1) in which sediment from land drainage i s reta ined. Almost a l l mangroves on the west coast are disturbed by man to a greater or l e s s e r degree, and therefore , well marked eco log ica l zonation i s often not seen. According t o Watson's (1928) innundation -c lasses described for Malaysian mangroves (Fig . 4 ) , the following eco log ica l zonation could be expected: Avicennia marina/ Sonneratia .alba/ Rhizophora spp. at the lower edge followed by Aegiceras corniculatum/ Avicennia off ic inal is /Bruguieta- spp./Ceriops tagal/Xylocarpus granatum followed by Acanthus i l i c i f o l i u s / Excoecaria agallocha/Lumnitzera racemosa/Sonneratia caseo lar i s followed by the mangrove a s soc ia t e s in the back mangrove. Lack of c l ear eco log ica l zonation in the mangroves of Sri Lanka may be part ly due to the small t i d a l amplitude experienced. However, whatever eco log ica l zonation that ex i s ted has great ly been disturbed by man. I t i s well known that once the w e l l - es tabl i shed eco log ica l balance i s disturbed, i t takes a long time for the proper zonation t o r e - e s t a b l i s h . If the disturbance per s i s t ed , then the proper zonation w i l l not r e ? e s tab l i sh at a l l . Development of many back mangrove spec ies such as Acrostichum aureum among the cere species even in the lower border of the mangrove, and the absence of tall trees indicate that the dominant trees had been removed. Tansley & Fritsch (1905) even at the beginning of t h i s century noted the e f f e c t of human interference on the Sri Lankan mangroves. K.H.G.M. OE SILVA AND S. BALASUBRAMANIAM 31 The wide distribution of Acanthus ilicifolius in the islands of the Negombo estuary (Fig. 2) seems to have led Abeywickrama (1964) to postulate that the earliest phanerogamic colonizer in mudflats is A.ilicifolius. According to him, this will first cover the entire surface of the small islands, and species such as Bruguiera spp. and Aegiceras corniculatum will invade later and form a mixed mangrove. Pinto (1982) also supports this hypothesis and suggests that the succession of plants in the islands of Negombo estuary took place with A^ ilicifolius pioneering followed by Rhizophora spp./B. gymnorhiza mixed vegetation and finally by Rhizophora spp./Avicennia marina vegetation. According to this view, at least one island, which is at present almost covered with A? ilicifolius, is still at the first stage of succession. However, this is unlikely as the islands are not of recent origin and since there is also no evidence that the islands had been totally denuded of all vegetation recently. It is more likely that A.ilicifolius has become the dominant plant as a result of selective destruction of other mangrove species by man allowing A^ ilicifolius to spread. In the mudflats exposed even today by the denudation of the vegetation, seedling establishment of Rhizophora spp. and Avicennia marina could be-observed, in this connection, it is noteworthy that Macintosh (1982) in his study of Malayan mangroves showed that in the Indo-Pacific region the pioneering mangrove species in coastal deposition shores are Avicennia spp. and Sonneratia spp. and those in more estuarine localities are Rhizophora spp. In the south, west and east coasts considerable tracts of mangroves have recently been cleared for construction of hotels and provision of recreation facilities to tourists. The diversion of the Mahaweli Ganga, the largest river in the country, for irrigation purposes may affect the mangroves in Trincomalee area (Fig. 1) by reducing the amount of nutrients carried to the estuary and also by carrying and discharging agricultural pesticides and residues. This scheme may also affect the mangroves in Kala Oya (west coast) and Maduru Oya (east coast) estuaries as water from the Mahaweli is diverted into these two rivers as well. The clearing of mangroves and the reclamation of mangrove land for various purposes go on, and no serious attempt has still been made to educate the local people about the value of the mangrove ecosystem. Without such education, legislation against the destruction of the mangroves will be of little use. Unless serious steps are taken urgently to conserve at least what is left, Sri Lankan mangroves are doomed. 32 MANGROVES OF SRI LANKA ACKNOWLEDGEMENTS We wish to thank Professor M.D. Dassanayake of Department of Botany and Dr.(Mrs.)P. Kumari de Silva of Department of Zoology, University of Peradeniya for their help and advice during this study. We are also grateful to Mr. N.M. Moiyadeen, O.I.C., NARA Regional Research Station, Kalpitiya for the help and facilities provided during the study, - and to Dr. J.M.R.S. Bandara, Head, Department of Agricultural Biology, and Professor K. Jayasena, Head, Department of Pharraocology, University of Peradeniya for providing vehicles to visit various sites. Thanxs are also due to Mr. T.S.B. Alagoda for drawing the figures. This work was supported by the research grant RGB/Gen/50 of the Natural Resources, Energy and Science Authority of Sri Lanka and we wish to record our appreciation to Dr. R.P. Jayawardene, Director General, and Dr. S. Liyanage, Assistant Director, for provision of funds. REFERENCES Abeywickrama, B.A. (1964). The estuarine vegetation of Ceylon. Proceedings of the Decca Symposium, UNESCO. 207-210. Amarasinghe, M.D. and Perera, W.K.T. (1984). A preliminary survey of peripheral vegetation communities of Puttalam lagoon and Dutch bay. Proceedings of the 40th Annual Session - Sri Lanka Ass.Adv.Sci., December 1984 (Abstract). Arulchelvam, K. (1968). Mangroves. Ceylon Forester, VIII (3 ? 4 ) : 1-34. Balasubramaniam, S. (1985). Tree flora in Sri Lanka. In: Proceedings of the International Conference on Timber Technology, . 24-26 October, 1984. Jayatilleke, A. (ed.) Univ. of Moratuwa, Sri Lanka. 58-67. Balasubramaniam, S., Rasiah:, P. and Ratnapala,M.L. (1970). Mangrove vegetation in Vadamarachchi lagoon. Phyta, 1: 15-20. 3 3 K.H.G.M. DE SILVA AND S. BALASUBRAMANIAM Jayawardene, R.P. (1985). The mangroves of Sri Lanka. Bakawan, IV (1): 8-10. Kanakaratne, M.D., Perera, W.R.T. and Fernando, B.U.S. (1984). An attempt at determining the mangrove coverage around Puttalam lagoon in Sri Lanka through remote sensing techniques. Proceedings of the 4th Asian Conference on Remote Sensing, Colombo. 1983. 9 PP. Macintosh, D.J. (1982). Fisheries and aquaculture significance of mangrove swamps, with special reference to Indo-West Pacific region. In: Recent Advances in Aquaculture. Muir, J.F. and Roberts, R.J. (eds.), Croom Helm, London. 3-85. Macnae, W (1968). A general account of the fauna and flora of mangrove swamps and forests in the Indo-West Pacific region. Adv. mar. Biol., 6: 73-270. Mepham, R.H. and Mepham, J.S. (1985). The flora of tidal forests - a rationalization of the use of the term 'mangrove'. S. Afr. J. Bot. 51 (2): 77-99. Odum, E.P. (1971). Fundamentals of ecology. W.B. Saunders Co., London. 352 p. Pinto, M.L. (1982). Distribution and zonation of mangroves in the northern part of the Negombo lagoon (Sri Lanka). J. natn. Sci. Coun. Sri Lanka, 10 (2): 245-255. Pritchard, D.W. (1967). what is an estuary: physical viewpoint. In: Estuaries, (Lauff, G.H. (ed.). Amer. Assoc. Adv. Sci. Publ. No. 83, Washington, D.C. 3- 5 PP. Tansley, A.G. and Fritsch, F.E. (1905). Sketches of vegetation at home and abroad. I. - The Flora of the Ceylon littoral. The New Phytologist, IV (1): 1-45. Tennent, E. (1859). Ceylon, an account of the island. Vol. I. Longman and Roberts, London. 86 p. Watson, J.G. (1928). Mangrove forests of the Malay peninsula. Malay For. Rec. 6: 1-275. TABLE 1 DISTRIBUTION OF MAJOR CORE SPECIES IN SOME OF THE MANGROVES IN WEST COAST OF SRI LANKA ABUNDANCE OF EACH SPECIES IS INDICATED IN AN ARBITRARY SCALE OF 1-5. (1 very common; W E T RIVERINE Gin Ganga Acanthus ilicifolius Aegiceiras corniculatum Avicennia marina Bruguiera cylindrica 5 i gyronorhiaa sexangula Ceriops tagal Excoecaria i agallocha Lumnitzera racemosa N^ga fruticana Rhizophora apiculata R. mucronata Sonneratia alba 4 2 4 4 4 3 common; frequent; Z O N E ESTUARINE Hikkaduwa estuary/ islets in Balapitiya estuary 2 3 4 3 4 3 Islets in Negombo estuary rare; 5 - very rare) D R Y Z O N E RIVERINE Kala Oya Pomparippu Aru S. caseolaris 1 4 3 1 2 4 5 2 3 5 3 2 3 2 3 3 5 (Maha Oya river estuary) 3 3 1 1 4 4 ESTUARINE Islets and mainland in Kalpitiya area 5 3 1 Mainland in Mundel estuary 2 2 3 4 1 3 5 K.H.G.M. DE SILVA ANO S. BALASUBRAMANIAM 35 TABLE 2 SPECIES OF MANGROVE PLANTS, MANGROVE ASSOCIATES, ANO SEA6RASSES OF SRI LANKA Core species of mangroves Acanthaceae Acanthus i l ic i fol ius L. Katu Ikiri (S) Kadalmulll (T) Avicenniaceae Avicennia marina (Forsk.) Vierh Manda (S) A. officinalis L. Combretaceae LumAitzera littorea (Jack; J.O. Volgt L^ racemosa Willd. Meliaceae Xylocarpus granatum Koenig X. moluccensis (Lamx)M. Roem Myrsinaceae ! icera lanco Aegi s corniculatum (L.) Rath beriya (S) Beriya (S) Mutti kadol (S) Mudu-delun (S) Avari kadol (S) Gin-pol (S) Mai kadol (S) Rath kadol (S) Ela kadol (S) Kanna (T) Venkandal (T) Kanna (T) Upu attha (T) Thlpparethai (T) Kontalai (T) Vettilaikanna (T) Narikandal (T) Palmae Nypa fruticans Wurmb. Rhizophoraceae Bruguiera cylindrica (L.) Blume B. gymnorrhiza (L.) Willd. B. sexangula (Lour.) Poir. Ceriops decandra (Griff.) Ding Hou Heen kadol (S) C tagal (Perr.) C.B. Rob. Heen kadol (S) Rhizophora apiculata Bl. Rana kadol (S) Rj. mucronata Lam. Murunga kadol (S) Rubiaceae Scyphiphora hydrophyllacea Gaertn.f. Sirukandal (T) Chlrukandal (T) Kandal (T) Kandal (T) 36 [A] K.H.G.M. OE SILVA AND S. BALASUBRAMANIAM Dalbergia candenatensis (Dennst.) Prain Derris heptaphy)la (L.)Merr. D. heterophylla (Willd.) Back (=0.ul1q1nosa) Ponqaroia pinnata (L.) Pierre Gramineae (Poaceae) Cyperus rotundus L. Zoysia matrella (L.) Merr. Guttiferae (Clusiaceae) Calophyllum inophyllum L. Lythraceae Pemphis acidula J.R. & G. Forst Malvaceae Hibiscus tiliaceus L. Thespesia populnea (L.) Solander ex Correa T. populneoides (Roxb.) Kosteletsky Hernandiaceae Hernandia oviqera L. Lecythidaceae Barrinqtonia asiatica (L.)Kurz B. racemosa (L.) Bl . Pandanaceae Pandanus odoratissimus L.F. Stercullaceae Heritiera littoralis Dryand Tamaricaceae Tamarix qallica L. Verbenaceae Clerodendron inerme (L.) Gaertn Premna inteqrifolia Lam (=P. foetida Reinw.) Kala-wel (S) Kalanduru (S) Domba (S) Belipatta (S) Gan suriya (S) Suriya (S) Diyamidella (S) Dlyamldella (S) Wetakeyya (S) Athuna (S) Wal-gurenda (S) Wal midi (S) Pungal (T) Korai (T) Punnal (T) Vellai (T) Poovarusu (T) Samuthram (T) Thall (T) Chomunthiri (T) Pichuvilathi (T) Erumai-mullai (T) 36[B] MANGROVES OF SRI LANKA Sonneratiaceae Sonneratla alba J. Sm. S. apetala Buch-Ham. S. caseolaris (L.) Engler Mangrove associates Annonaceae Annona glabra L. Apocynaceae Cerbera odollam Gaertn. Qchrosia oppositifolia (Lam) Scnum Ai zoaceae Sesuvlum portulacastrum (L.) L. Bignoniaceae Oolichandrone spathacea (L.f . ) K. Sehum. Chenopodi aceae Arthrocnemum Indicum (Willd.)Moq. Salicornia brachiata Roxb. Suaeda maritima (L.) Dumort Sj. monoica Forsk. ex J.F.Gmel S. nudiflora (Willd)Moq. Compositae (Asteraceae) Sphaeranthus amaranthoides Burmf. S^ africanus L. Cyperaceae ? Fimbristylis ferruginea (L.) Vahl Schoenoplectus Uttoralis (Schrad)Palla. Euphorbiaceae Excoecarla agallocha L. Fabaceae (Leguminosae) Cynometra 1ripa Kostel C. bijuga var. mimosoides Merr. Sudu mal klrala (S) Vellal-kinnai (T) Rath mal klrala (S) Klnnal (T) Wal anoda (S) Kadal-attha (T) Gon-kaduru (S) Nachchukal (T) D1ya-Danga (S) V1lpath1r1 (T) Mankulanchl (T) Kotanal (T) Um1r1 (T) Um1r1 (T) Um1r1 (T) Thelakeerlya (S) Th1lla1 (T) Kadupull (S) MANGROVES OF SRI LANKA 3 7 Healparasites on mangroves & mangrove associates Loranthaceae Dendrophthoe falcata (L.f .) Danser PI111a (S) Kuruvlchal (T) Aeglceras corniculatum Viscaceae Vlscum orlentale Willd. on Excoecaria agallocha and Tehspesla populnea Mangrove fern Adlantaceae Acrostichum aureum L. Karenkoku (S) M1nn1 (T) Sea grasses Hydrocharitaceae Enhalus acoroldes (L.f.) Royle Halophila deciplens Ostenf. H. oval1s (R.Br.) Hook f. Thalassia hemprichii (Ehrenb.) Aeschers Najadaceae Najas graminea Del. marina L. Potamogetonaceae Cymodocea serrulata (R. Br.) Aeschers & Magnus Halodule uninervis (Forsk.) Aeschers (aD1pianthera uninervis) (Forsk) Aeschers) Potamogeton pectinatus L. Ruppia maritlma L. Syrinqodium Isoetifolium (Aeschers) Dandy 38 K.H.G.M. DE SILVA AND S. BALASUBRAMANIAM LEGENDS TO FIGURES FIGURE 1: Inland water resources of Sri Lanka. Estuaries studied are] indicated by arrows. Dry zone and wet zone are also indicated. FIGURE 2. (a) upper part of Kalpitiya estuary and (b) upper part of Negombo estuary. Dotted are magroves. areas FIGURE 3: (a) Balapitiya estuary and (b) Hikkaduwa estuary and surrounding plantations. Dotted areas are mangroves. C- coconut; N-cinnamon; R-rubber; M-marsh. crop FIGURE 4: Ecological distribution of major mangrove core species according to Watson's (1928) innundation classes. Classes 1,2,3,4, and 5 indicate areas inundated by all high tides, medium high tides, normal high tides, spring tides and equinoctial tides respectively. INDIAN OCEAN E S T U A R Y PERIYA , A R C H C H A L K I R I M U N D E L S I N N A O O O A K A R E N T I V U ^ A R C H C H A L K O V J L K U D A H ^ K ^ N E R U M A T V U ^ff " " M A T T U T I V U ^ A M B A N T A T I V U PULLUPIDDI ? ESTUARY Q K A N O A K U D A H Inundation Classes Acanthus ilicifolius Aegiceras corniculatum Avicennia marina A. officinalis Bruguiera cylindrica B. gymnorhiza 8 . sexangula Ceriops tag a I Excoecaria agallocha Lumnitzera racemosa Nypa fruticans Rhizophora apiculata ft. mucronata Sonneratia alba S . caseolaris