Responses Of The Mangrove Ecosystem To . - Acpa.botany.pl
Acta Palaeobotanica 59(2): 391–409, 2019DOI: 10.2478/acpa-2019-0013e-ISSN 2082-0259ISSN 0001-6594Responses of the mangrove ecosystemto Holocene environmental change in theSundarban Biosphere Reserve, IndiaARGHYA KUMAR HAIT 1* and HERMANN BEHLING 212Department of Botany, City College, Kolkata – 700 009, India; e-mail: akhait@hotmail.comGeorg-August-University of Göttingen, Albrecht-von-Haller-Institute for Plant Sciences, Department ofPalynology and Climate Dynamics, Untere Karspüle 2, 37073 Göttingen, GermanyReceived 17 March 2019; accepted for publication 23 September 2019ABSTRACT. The Sundarban Mangrove Forest in the Sundarban Biosphere Reserve, located at the mouth ofthe Ganga–Brahmaputra Delta in India, is the most diverse mangrove ecosystem in the world. Sediment coreswere taken from two widely separated islands in that reserve: Chamta (CMT) and Sudhyanyakhali (SDK). Pollen analysis and radiocarbon dating were used to study the Holocene development and dynamics of this uniqueecosystem. Modern pollen rain study reveals a strong relation between modern pollen rain and the presentvegetation, as well as a high rate of Phoenix palludosa pollen production.The pollen records indicate that mangrove existed at CMT from 5960 and at SDK from 1520 cal yr BP. Changes in relative sea level, includingthe frequency and intensity of inundation as well as fluctuating precipitation, have been the major factors alongwith geomorphic processes that control the development and dynamics of the mangrove in the area during theHolocene. The mid Holocene mangrove at CMT declined, to be progressively replaced by successive communities, and eventually reached climax stage, while the SDK site is transitional in nature. The mangrove respondsrapidly to changes in environmental conditions at both locations. Because of large-scale anthropogenic interventions, it is unlikely that similar rapid responses will occur in the future.KEYWORDS: Delta, Holocene vegetation dynamics, pollen analysis, global change, sea levelINTRODUCTIONThe Indian Sundarbans, located at 21 32′–22 40′N and 88 5′–89 10′E in the state of WestBengal, are famous for their extensive mangrove vegetation (Fig. 1) and are a BiosphereReserve. The total area of the Indian Sundarbans is 9630 km2, of which 4264 km2 is covered by mangrove, and 1738 km2 by creeks,creeklets, estuaries and other waterways. Thearea has about 124 low-lying swampy islandsformed by an intricate network of differenttypes of water channels, 84 of which are inhabited by mangroves. This forest is unique as it isthe most diverse mangrove ecosystem (Sanyal1996, 2001a, Gopal & Chauhan 2006) and oneof the two mangrove tiger lands in the world.*Corresponding authorThe Sundarban mangrove belongs to the category of littoral and swamp forest accordingto the vegetation mapping of the entire region(Das Gupta 1975). This littoral and swampforest has been surveyed for vegetation types,plant distribution, plant associations, ecological succession (Rao & Sastry 1974, Chanda& Datta 1986, Naskar & Guha Bakshi 1989,Chaudhuri & Choudhury 1994, Blasco et al.1996, Debnath & Naskar 1999, Naskar & Mandal 1999, Ellison et al. 2000, Gopal & Chauhan 2006, Giri et al. 2014, Ghosh et al. 2015,Barik et al. 2017, Danda et al. 2017, Mandalet al. 2019), modern pollen distribution (Pandey & Holt 2018, Pandey & Minckley 2019),change detection (Ghosh et al. 2015, Ghoshet al. 2016), impact of climate change and sea
392A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019Fig. 1. Location maplevel fluctuations on mangrove (Raha et al.2012, Ellison 2015, Alongi 2015, Ward et al.2016), biogeochemical and other studies (Ramanathan et al. 2009, Manna et al. 2010, Donatoet al. 2011, Chowdhury & Maiti 2016, Majumder et al. 2016, Dutta et al. 2017).Recent reviews by Sanyal (2001a) indicatethe presence of 18 major mangrove specieswhich are exclusively found in the inter-tidalzone and manifest typical mangrove adaptations such as pneumatophores, vivipary andsalt glands. Fifteen other mangrove species(also found exclusively in the inter-tidal zone)and 51 species of back mangrove are foundin the inter-tidal as well as beyond the tidalzone. The important tree taxa of the presentday Sundarban forest are Avicennia sp., Excoecaria sp., Sonneratia sp., Rhizophora sp., Aegiceras sp., Bruguiera sp., Ceriops sp., Heritierasp., Kandelia sp. and Xylocarpus sp. The mangrove palm Nypa sp. and Phoenix sp., herbslike Acanthus sp., Suaeda sp. and Porteresiasp., and the mangrove fern Acrostichum sp.are other important mangrove taxa present inthe Sundarban forest.This estuarine ecosystem has environmental, ecological and economic significance to therapidly growing population of this region. Theforest acts as a buffer and saves the hinterlandfrom cyclones and storms during the monsoon,retaining sediment and thereby retardingcoastal erosion. The ecosystem is very rich innutrients and provides a habitat and breedinggrounds for many species of fish and other economically important organisms. A large section of the population is also directly dependent on this forest for its livelihood (Chaudhuri& Choudhury 1994, Broadus 1996, Stanley& Hait 2000a, Gopal & Chauhan 2006).Imbalances in this ecosystem have serioussocio-economic and ecological implications.The delta was affected during the early andmid Holocene by natural factors such as aninclination towards the east and rising sealevel (Blasco et al. 1996, Stanley & Hait 2000a,Gopal & Chauhan 2006, Hait & Behling 2009,
393A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019Barui 2011, Das 2014, Mishra et al. 2016,Sarkar & Sen 2019). In recent times, increasinghuman activity in the form of land reclamationand withdrawal of river water upstream hasbeen responsible for habitat loss and enhancedsalinity of both the water and soil (Millimanet al. 1989, Nazrul-Islam 1993, Alam 1996,Blasco et al. 1996, Allison 1998, Stanley & Hait2000a, Gopal & Chauhan 2006, Zaman et al.2013, Ghosh et al. 2015). Studies also indicatethe presence of toxic material of anthropogenicorigin in the soil, water and air of the Sundarban mangrove ecosystem (Santra 1994, Sikdar& Hait 1997, Sikdar et al. 1998, Saha et al.2005, Gopal & Chauhan 2006, Manna et al.2010, Ghosh et al. 2015, Chowdhury & Maiti2016). Conservation of this unique ecosystem,which provides many benefits, is essential.There have been many key advances in palaeobiological methodologies which may greatlyimprove the quality and relevance of palaeoenvironmental and palaeoecological studies forconservation biology. Palaeobiological studies can address questions about the biologicalvulnerability and resilience of a single taxon,a community, or a whole ecosystem facingenvironmental changes. This relates directlyto conservation biology, including the design ofbiological reserves, the effect of biological andenvironmental change on ecosystem services,the stability of biogeochemical cycles, and thedirect and indirect impacts of invasive species and extinction of species (Flessa & Jackson 2005, Willis & Birks 2006, Dietl & Flessa2009, 2011, Jackson & Hobbs 2009, Willis et al.2010, Birks 2012, Setyaningsih et al. 2019).This study was undertaken to understandthe biological, ecological and environmentalchanges that have taken place in and aroundstudy sites located in the core and buffer areaof the Sundarban Biosphere Reserve, India,during the Holocene. This paper reports thefirst record of the Holocene vegetation andenvironmental history of the core and bufferarea. The information obtained through thisstudy can be used to formulate realistic conservation and management plans for this biologically unique ecosystem.Another objective was to understand therelationship between the extant mangrove vegetation and modern pollen rain data, for betterinterpretation of fossil pollen data in studies ofHolocene vegetation and palaeoenvironmentalreconstruction.STUDY AREALOCATIONThe study sites are located inside the deltaicSundarban mangrove forest (Fig. 1), which isa Biosphere Reserve. The forest forms the southwestern part of the large Ganga–BrahmaputraDelta, between the Hugli River to the west andHaribhanga River to the east. The northernlimit is demarcated by the Dampier–HodgesLine. The reserve is divided into core and bufferareas for management and conservation purposes. No human activity is allowed inside thecore area. The area is further demarcated intothree Wildlife Sanctuaries (Sajnekhali, Haliday,Lothian) and one National Park. The entire areaof the National Park and the Sajnekhali Wildlife Sanctuary are included in the Project TigerArea. This whole Biosphere Reserve has 22 forest blocks, of which the Sudhyanyakhali (SDK)forest campsite of the Pirkhali forest block inthe buffer area and the Chamta (CMT) forestcampsite of the Chamta forest block in the corearea of the Biosphere Reserve were selected forsediment coring.GEOLOGY AND GEOMORPHOLOGYThe Ganga–Brahmaputra delta representsa late Quaternary sedimentation history (Stanley & Hait 2000a, Sarkar et al. 2009) and isunderlain as well as bounded on three sides(east, north and west) by Precambrian metamorphic and Gondwana sediments and Mesozoic trap volcanics (Chanda et al. 1999). It isopen to the sea on the south. Late Quaternaryneotectonic activity that induced tilting of thedelta region to the east altered the course of theGanga–Brahmaputra–Meghna River system,resulting in an eastward shift in delta progradation (Sanyal 1990, Chaudhuri & Choudhury1994, Blasco et al. 1996, Stanley & Hait 2000a).The general lithology of the Holocene deposits of the coastal areas comprises fine to mediumsand, silt and clay. Peat layers are also foundin the north and northwest of the areas outside the present-day mangrove forest (Stanley& Hait 2000a, Stanley et al. 2000). The elevation is gentle, and most of the area is not morethan 1 m above the present-day mean sea level(Delft Hydraulics 1989, Milliman et al. 1989,Jelgersma 1994, Broadus 1996).Geomorphologically, the area is a set of tidedominated estuaries with numerous linear tidal
394A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019sand bars and an extensive network of tidalchannels, of which the larger channels (often1.5 to 20 km wide) run north–south (Gopal& Chauhan 2006). The mouth of the estuary isfunnel-shaped because of the high tidal amplitude of 3.5 to 5.0 m (Chaudhuri & Choudhury1994). A variety of habitats occur, includingbeaches, estuaries, swamps, tidal flats, tidalcreeks, creek lets, coastal dunes and backdunes, and many of these are covered by densemangrove (Sanyal & Ball 1986, Paul 1987a, b,Naskar & Guha Bakshi 1989, Gopal & Chauhan 2006).88 19′E) within the Reserve, which is an isolated island. The site has profuse growth ofAvicennia sp. Ceriops decandra, Bruguieragymnorrhiza and Excoecaria agallocha arefound scattered in this block, with very lowrepresentation of other mangrove taxa. Thethird site (Bakkhali [E] 21 37′N, 88 18′E) isclose to the coast; there, the predominant vegetation is Excoecaria agallocha, followed byAegiceras corniculatum and Phoenix paludosa.STUDY SITESFIELD SAMPLINGTwo different areas within the mangroveforest were selected for subsurface sedimentsampling (Fig. 1). The first site, Sudhyanyakhali (SDK [A]: 22 06′09″N, 88 54′50″E) in thePirkhali forest block, is located towards thenorth of the forest in the buffer area, and is partof the Sajnekhali Wildlife Sanctuary. The corewas taken from an area which is apparentlystable and not inundated regularly by tides.Ceriops decandra, Excoecaria agallocha andXylocarpus mekongensis are the most commonmangrove taxa in this block. Bruguiera gymnorhiza and Avicennia sp. are common alongthe creeks and creeklets. This forest block hasa thick undergrowth of Phoenix paludosa.The second site, Chamta (CMT [B]:21 51′40″N, 88 54′56″E) of the Chamta forestblock, is located in the core area of the Reserve,belongs to the category of National Park, andis remote and inaccessible during most timesof the year. The block is characterized by highrepresentation of low dense stands of Ceriopsdecandra, followed by Excoecaria agallocha.Species of Xylocarpus are also found scatteredin this block. Other mangrove species are alsopresent but in very low numbers. The corewas collected from an area of Ceriops stands 300 meters from the nearest active channel,which is not regularly flooded by tidal water.Pollen traps were installed for collectionof modern pollen rain data in three differentareas in the buffer area of the Reserve (Fig. 1).The first site (Jharkhali [C] 22 00′N, 88 42′E)is towards the western part of the Reserve,where the vegetation is heterogeneous, withpredominance of Avicennia, followed byExcoecaria. The second site is in the middleof a wildlife sanctuary (Lothian [D] 21 38′N,MATERIALS AND METHODSSediment samples were collected from two differentareas inside the Sundarban mangrove forest (Fig. 1),using a Russian sampler. Both cores (SDK, CMT) are300 cm in length. Lithological descriptions were madeand subsampling for palynological analysis was doneimmediately after recovery of the core.Modern pollen rain data were collected with thetype of pollen traps used by Behling et al. (2001) fromthree different locations (C, D & E; Fig. 1). Trapping ofairborne pollen grains and spores was done using plastic tubes (h 11.5 cm, diam. 2.7 cm) filled with 5 cm3 glycerin and a few drops of phenol to avoid fungal growth.The tubes were covered with nylon mesh to avoidinsects and debris. A set of five traps was installed 15 m apart in each location. In the area inhabited byPhoenix paludosa, which is less inundated, the trapswere installed 50 cm above the surface. In the swampymangrove areas the traps were installed at 2 m heightto avoid tidal inundation. The collection period lastedone year from September 2002 to August 2003.POLLEN ANALYSISFor palynological study, 0.5 cm3 sediment was takenat 5 cm intervals along the cores. Prior to processingof the sediment and pollen rain samples, one tablet ofexotic Lycopodium spores was added for calculationof pollen concentration (grains/cm3) and pollen accumulation rate (grains/cm2/yr). Palynological preparations were made following standard pollen analyticaltechniques (Erdtman 1969, Faegri & Iverson 1989).Slides were prepared with sample residues mountedin glycerin jelly medium. Published pollen morphological descriptions were consulted for identificationof pollen grains and spores (Thanikaimoni 1987). Thereference collection of the second author was also consulted for identification purposes. On the basis of ecological requirements and life form, taxa were groupedin the following categories: mangrove, non-mangrove(NM), herb, hypersaline herb (HSB), aquatics, fern,mangrove fern (MF), fungi, marine. Most of the samples were counted to a minimum of 300 pollen grains,including arboreal and nonarboreal pollen. This pollensum excludes aquatic pollen, fern and fungal sporesand marine taxa. The palynological data are presentedin pollen diagrams as percentages of the pollen sum.
395A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019TILIA was used for calculations and CONISS was usedfor cluster analysis of pollen taxa. Cluster analysis isbased on stratigraphically constrained pollen percentage data included in the pollen sum. TGVIEW was usedto plot the pollen diagrams (Grimm 1987). The zonationof both cores is based on the cluster analysis and onconspicuous changes in the pollen assemblages.RESULTSSTRATIGRAPHYThe general lithological unit in both cores issilt and clay. No peat was recorded in any ofthe cores. The lower part of the Chamta (CMT)core is black, sticky clay (300–290 cm). Thenext part of the core (290–0 cm) is grey siltyclay with frequent occurrence of decomposedwood fragments.The lower part (300–190 cm) of the Sudhyanyakhali (SDK) core is grey silty clay;a decomposed wood fragment was noted in the300–230 cm interval of this sequence. The middle section (190–90 cm) is grey sticky clay. Thelithology changes from clayey silt (90–45 cm) tosilty clay (45–0 cm) in the top of the core.CHRONOLOGYTwo organic-rich sediment samples (1 cmthick) from each location were taken and datedby accelerator mass spectrometry (AMS) at theLeibnitz laboratory of Christian–Albrecht–Universität in Kiel, Germany (Tab. 1). Onlyorganic bulk sediments could be taken for dating. The dates were calibrated using Cal Pal(Weninger et al. 2004).Two radiocarbon dates for the CMT coreindicate deposits of mid and late Holoceneage. The extrapolated age of the core base at300 cm of the CMT core is 6083 cal yr BP. Sediment accumulation seems to be continuous,Table 1. List of AMS radiocarbon dates of CMT and SDK coreLab. no.Depthcm14AgeC yr BPAgecal yr BPMaterialdatedLocation: CMTKIA233571032830 402935 51KIA233582955195 355955 : SDKKIA233601209960 60 11,404 125KIA233592951635 301519 40Organic-richsedimentOrganic-richsedimentprobably without any gaps until the top of thecore; this is probably modern in age.The SDK core is younger. It began to accumulate at 1520 cal yr BP (1635 30 14C yrBP) at 295 cm core depth. The radiocarbondate from the upper section of this core at120 cm shows a very old age of 9960 60 14Cyr BP. The record of an old age in a youngersequence in Holocene deltaic sediments probably is related to the remobilization of sediment that prevails in fluvial and deltaic plain(Stanley & Hait 2000b). To some extent thepossible contamination of the sequence witholder pollen and spores has to be taken intoaccount. This old radiocarbon date has beenexcluded from the age calculations of each pollen zone. The age calculations for the zones ofthe SDK core are therefore somewhat limitedand should be considered an approximation.DESCRIPTION OF THE MODERNPOLLEN RAIN DATATen pollen trap samples from three differentlocations (C–E) were used for the modern pollenrain study. Three other samples from locationC (Jharkhali) and two other samples from location E (Bakkhali) were contaminated, due totidal inundation. Palynological study revealed13 pollen types and 1 fern spore. Ten of the13 pollen types belonged to mangrove (Avicennia, Phoenix, Excoecaria, Acanthus, Aegiceras,Bruguiera, Ceriops, Rhizophora, Sonneratioaand Xylocarpus), 1 to non-mangrove (Borassus), 1 to a hypersaline herb (Suaeda) associated with the mangrove environment, and 1 toPoaceae. The only spore type was assignable tothe mangrove fern Acrostichum aureum. Thepercentages of the most frequent pollen taxaand the pollen influx rates are shown in Figure2. The sum of each group and the fern sporeAcrostichum aureum are also included in thediagram.The three locations are inhabited by different types of mangrove plants. Locations C andD have higher representation of Avicennia,while Phoenix is dominant at location E, followed by Excoecaria and Aegiceras. Non-mangrove tree taxa are almost absent except in onesample at location C. Herb pollen are commonin most of the locations except E. The dominance of Avicennia in the modern pollen rainis directly related to the higher representationof this plant in locations C and D. Regardingthe domination of Phoenix at location E, we
Fig. 2. Modern pollen rain data of Jharkhali, Lothian and Bakkhali in the Sundarbans396A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019
A.K. Hait and H. Behling / Acta Palaeobotanica 59(2): 391–409, 2019note that Phoenix is a high producer of pollen,while Avicennia, Excoecaria and Aegiceras arerelatively low producers.Pollen influx shows marked variationbetween sites C–D and E. It is very high (avg.20 200 grains/cm2/yr) for the Excoecaria /Phoenix-dominated forest at location E and very lowfor the Avicennia-dominated mixed mangroveforest at locations C and D (2450 grains/cm2/yr).Description of the Chamta (CMT)Pollen DiagramThe whole CMT pollen record is marked byabundant (80–90%) poll
1 Department of Botany, City College, Kolkata – 700 009, India; e-mail: akhait@hotmail.com 2 Georg-August-University of Göttingen, Albrecht-von-Haller-Institute for Plant Sciences, Department of Palynology and Climate Dynamics, Untere Karspüle 2, 37073 Göttingen, Germany Received 17 March 2019; accepted for publication 23 September 2019 ABSTRACT. The Sundarban Mangrove Forest in the .
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