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Running Head 'Biology of Mangroves' BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS 1 Biology of Mangroves and Mangrove Ecosystems ADVANCES IN MARINE BIOLOGY VOL 40: 81-251 (2001) K. Kathiresan1 and B.L. Bingham2 1Centre of Advanced Study in Marine Biology, Annamalai University, Parangipettai 608 502, India 2Huxley College of Environmental Studies, Western Washington University, Bellingham, WA 98225, USA e-mail [email protected] (correponding author) 1. Introduction.............................................................................................. 4 1.1. Preface........................................................................................ 4 1.2. Definition ................................................................................... 5 1.3. Global distribution ..................................................................... 5 2. History and Evolution ............................................................................. 10 2.1. Historical background ................................................................ 10 2.2. Evolution.................................................................................... 11 3. Biology of mangroves 3.1. Taxonomy and genetics.............................................................. 12 3.2. Anatomy..................................................................................... 15 3.3. Physiology ................................................................................. 18 3.4. Biochemistry ............................................................................. 20 3.5. Pollination biology..................................................................... 21 3.6. Reproduction, dispersal and establishment................................ 22 3.7. Biomass and litter production .................................................... 24 4. Mangrove-associated flora 4.1. Bacteria ...................................................................................... 27 4.2. Fungi and fungus-like protists.................................................... 29 4.3. Microalgae.................................................................................. 33 4.4. Macroalgae................................................................................. 34 4.5. Seagrasses .................................................................................. 36 4.6. Saltmarsh and other flora ........................................................... 37 5. Mangrove-associated fauna 5.1. Zooplankton ............................................................................... 38 5.2. Sponges and Ascidians............................................................... 39 5.3. Epibenthos, infauna, and meiofauna .......................................... 41 5.4. Prawns, shrimp and other crustaceans ....................................... 43 5.5. Crabs .......................................................................................... 45 5.6. Insects......................................................................................... 49 5.7. Mollusks..................................................................................... 50 5.8. Fish............................................................................................. 52 BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS 2 5.9. Amphibians and Reptiles ........................................................... 56 5.10. Birds ......................................................................................... 56 5.11. Mammals.................................................................................. 57 6. Responses of mangroves and mangrove ecosystems to stress................ 58 6.1. Responses to light ...................................................................... 58 6.2. Responses to gases ..................................................................... 59 6.3. Responses to wind...................................................................... 61 6.4. Responses to coastal changes..................................................... 61 6.5. Responses to tidal gradients and zonation ................................. 63 6.6. Responses to soil conditions ...................................................... 64 6.7. Responses to salinity.................................................................. 66 6.8. Responses to metal pollution ..................................................... 67 6.9. Responses to organic pollution .................................................. 69 6.10. Responses to oil pollution........................................................ 70 6.11. Responses to pests.................................................................... 71 6.12. Responses to anthropogenic stress........................................... 73 6.13. Responses to global changes.................................................... 75 7. Ecological role of mangrove ecosystems 7.1. Litter decomposition and nutrient enrichment ........................... 76 7.2. Food webs and energy fluxes..................................................... 78 8. Concluding remarks ................................................................................ 80 Acknowledgements........................................................................... 82 References......................................................................................... Mangroves are woody plants that grow at the interface between land and sea in tropical and sub-tropical latitudes where they exist in conditions of high salinity, extreme tides, strong winds, high temperatures and muddy, anaerobic soils. There may be no other group of plants with such highly developed morphological and physiological adaptations to extreme conditions. Because of their environment, mangroves are necessarily tolerant of high salt levels and have mechanisms to take up water despite strong osmotic potentials. Some also take up salts, but excrete them through specialized glands in the leaves. Others transfer salts into senescent leaves or store them in the bark or the wood. Still others simply become increasingly conservative in their water use as water salinity increases. Morphological specializations include profuse lateral roots that anchor the trees in the loose sediments, exposed aerial roots for gas exchange and viviparous water-dispersed propagules. Mangroves create unique ecological environments that host rich assemblages of species. The muddy or sandy sediments of the mangal are home to a variety of epibenthic, infaunal, and meiofaunal invertebrates. Channels within the mangal support communities of phytoplankton, zooplankton, and fish. The mangal may play a special role as nursery habitat for juveniles of fish whose adults occupy other habitats (e.g., coral reefs and seagrass beds) Because they are surrounded by loose sediments, the submerged mangroves roots, trunks, and branches are islands of habitat that may attract rich epifaunal communities including bacteria, fungi, macroalgae, and invertebrates. The aerial roots, trunks, leaves BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS 3 and branches host other groups of organisms. A number of crab species live among the roots, on the trunks or even forage in the canopy. Insects, reptiles, amphibians, birds and mammals thrive in the habitat and contribute to its unique character. Living at the interface between land and sea, mangroves are well adapted to deal with natural stressors (e.g., temperature, salinity, anoxia, UV). However, because they live close to their tolerance limits, they may be particularly sensitive to disturbances like those created by human activities. Because of their proximity to population centers, mangals have historically been favored sites for sewage disposal. Industrial effluents have contributed to heavy metal contamination in the sediments. Oil from spills and from petroleum production has flowed into many mangals. These insults have had significant negative effects on the mangroves. Habitat destruction through human encroachment has been the primary cause of mangrove loss. Diversion of freshwater for irrigation and land reclamation has destroyed extensive mangrove forests. In the past several decades, numerous tracts of mangrove have been converted for aquaculture, fundamentally altering the nature of the habitat. Measurements reveal alarming levels of mangrove destruction. Some estimates put global loss rates at one million ha y-1, with mangroves in some regions in danger of complete collapse. Heavy historical exploitation of mangroves has left many remaining habitats severely damaged. These impacts are likely to continue, and worsen, as human populations expand further into the mangals. In regions where mangrove removal has produced significant environmental problems, efforts are underway to launch mangrove agroforestry and agriculture projects. Mangrove systems require intensive care to save threatened areas. So far, conservation and management efforts lag behind the destruction; there is still much to learn about proper management and sustainable harvesting of mangrove forests. Mangroves have enormous ecological value. They protect and stabilize coastlines, enrich coastal waters, yield commercial forest products and support coastal fisheries. Mangrove forests are among the world’s most productive ecosystems, producing organic carbon well in excess of the ecosystem requirements and contributing significantly to the global carbon cycle. Extracts from mangroves and mangrove-dependent species have proven activity against human, animal and plant
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