Mangrove forest: Difference between revisions

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File:Great egrets, Ardea alba, fishing.jpg|Three [[great egret]]s fishing along a mangrove shore
File:Great egrets, Ardea alba, fishing.jpg|Three [[great egret]]s fishing along a mangrove shore
File:Pelicans and double breasted cormorants in the mangroves.jpg|Pelicans and cormorants in the mangroves
File:Pelicans and double breasted cormorants in the mangroves.jpg|Pelicans and cormorants in the mangroves
</gallery>

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====Mangrove crabs====
Mangrove forests are among the most productive and diverse ecosystems on the planet, despite limited nitrogen (N) availability. Under such conditions, animal-microbe associations ([[holobiont]]s) are often key to ecosystem functioning. Here, we investigated the role of [[fiddler crab]]s and their [[carapace]]-associated [[microbial biofilm]] as hotspots of microbial N transformations and sources of N within the mangrove ecosystem.<ref name=Zilius2020 />

Among coastal ecosystems, mangrove forests are of great importance as they account for three quarters of the tropical coastline and provide different ecosystem services.<ref>{{cite journal |doi = 10.1111/geb.12155|title = Ecological role and services of tropical mangrove ecosystems: A reassessment|year = 2014|last1 = Lee|first1 = Shing Yip|last2 = Primavera|first2 = Jurgene H.|last3 = Dahdouh-Guebas|first3 = Farid|last4 = McKee|first4 = Karen|last5 = Bosire|first5 = Jared O.|last6 = Cannicci|first6 = Stefano|last7 = Diele|first7 = Karen|last8 = Fromard|first8 = Francois|last9 = Koedam|first9 = Nico|last10 = Marchand|first10 = Cyril|last11 = Mendelssohn|first11 = Irving|last12 = Mukherjee|first12 = Nibedita|last13 = Record|first13 = Sydne|journal = Global Ecology and Biogeography|volume = 23|issue = 7|pages = 726–743|hdl = 10862/2247}}</ref><ref>{{cite book |doi = 10.1016/S0065-2881(01)40003-4|title = Biology of mangroves and mangrove Ecosystems|series = Advances in Marine Biology|year = 2001|last1 = Kathiresan|first1 = K.|last2 = Bingham|first2 = B.L.|volume = 40|pages = 81–251|isbn = 9780120261406}}</ref> Mangrove ecosystems generally act as a net sink of carbon, although they release organic matter to the sea in the form of dissolved refractory macromolecules, leaves, branches and other debris.<ref name=Dittmar2006>{{cite journal |doi = 10.1029/2005GB002570|title = Mangroves, a major source of dissolved organic carbon to the oceans|year = 2006|last1 = Dittmar|first1 = Thorsten|last2 = Hertkorn|first2 = Norbert|last3 = Kattner|first3 = Gerhard|last4 = Lara|first4 = Rubén J.|journal = Global Biogeochemical Cycles|volume = 20|issue = 1|pages = n/a|bibcode = 2006GBioC..20.1012D}}</ref><ref>{{cite journal |doi = 10.1016/j.aquabot.2007.12.005|title = Organic carbon dynamics in mangrove ecosystems: A review|year = 2008|last1 = Kristensen|first1 = Erik|last2 = Bouillon|first2 = Steven|last3 = Dittmar|first3 = Thorsten|last4 = Marchand|first4 = Cyril|journal = Aquatic Botany|volume = 89|issue = 2|pages = 201–219|url = https://lirias.kuleuven.be/handle/123456789/203718}}</ref> In pristine environments, mangroves are among the most productive ecosystems on the planet, despite growing in tropical waters that are often nutrient depleted.<ref>{{cite journal |doi = 10.1093/treephys/tpq048|title = Nutrition of mangroves|year = 2010|last1 = Reef|first1 = R.|last2 = Feller|first2 = I. C.|last3 = Lovelock|first3 = C. E.|journal = Tree Physiology|volume = 30|issue = 9|pages = 1148–1160|pmid = 20566581}}</ref> The refractory nature of the organic matter produced and retained in mangroves can slow the recycling of nutrients, particularly of nitrogen (N).<ref name=Dittmar2006 /><ref>{{cite journal |doi = 10.1007/BF01204455|title = Sedimentary C/S relationships in a large tropical estuary: Evidence for refractory carbon inputs from mangroves|year = 1995|last1 = Woolfe|first1 = Ken J.|last2 = Dale|first2 = Paul J.|last3 = Brunskill|first3 = Gregg J.|journal = Geo-Marine Letters|volume = 15|issue = 3–4|pages = 140–144|bibcode = 1995GML....15..140W|s2cid = 128709551}}</ref> Nitrogen limitation in such systems may be overcome by microbial dinitrogen (N2) fixation when combined with high rates of bioturbation by macrofauna.<ref>{{cite journal |doi = 10.1023/A:1005850032254|year = 1997|last1 = Woitchik|first1 = A.F.|last2 = Ohowa|first2 = B.|last3 = Kazungu|first3 = J.M.|last4 = Rao|first4 = R.G.|last5 = Goeyens|first5 = L.|last6 = Dehairs|first6 = F.|journal = Biogeochemistry|volume = 39|pages = 15–35|s2cid = 91314553}}</ref><ref>{{cite journal |doi = 10.1128/aem.35.3.567-575.1978|title = Biological Dinitrogen Fixation (Acetylene Reduction) Associated with Florida Mangroves|year = 1978|last1 = Zuberer|first1 = D. A.|last2 = Silver|first2 = W. S.|journal = Applied and Environmental Microbiology|volume = 35|issue = 3|pages = 567–575|pmid = 637550|pmc = 242881|bibcode = 1978ApEnM..35..567Z}}</ref><ref name= Zilius2020 />

Bioturbation by macrofauna affect N availability and multiple N-related microbial processes through sediment reworking, burrow construction and bioirrigation, feeding and excretion.<ref>{{cite journal |doi = 10.3354/meps09506 |title = What is bioturbation? The need for a precise definition for fauna in aquatic sciences |year = 2012 |last1 = Kristensen |first1 = E. |last2 = Penha-Lopes |first2 = G. |last3 = Delefosse |first3 = M. |last4 = Valdemarsen |first4 = T. |last5 = Quintana |first5 = CO |last6 = Banta |first6 = GT |journal = Marine Ecology Progress Series |volume = 446 |pages = 285–302 |bibcode = 2012MEPS..446..285K }}</ref> Macrofauna mix old and fresh organic matter, extend oxic–anoxic sediment interfaces, increase the availability of energy-yielding electron acceptors and increase N turnover via direct excretion.<ref>{{cite journal |doi = 10.1080/0275754031000155474|title = It's a dirty job but someone has to do it: The role of marine benthic macrofauna in organic matter turnover and nutrient recycling to the water column|year = 2003|last1 = Welsh|first1 = David T.|journal = Chemistry and Ecology|volume = 19|issue = 5|pages = 321–342|s2cid = 94773926}}</ref><ref>{{cite journal |doi = 10.5194/bg-10-7829-2013 |title = Stimulation of microbial nitrogen cycling in aquatic ecosystems by benthic macrofauna: Mechanisms and environmental implications |year = 2013 |last1 = Stief |first1 = P. |journal = Biogeosciences |volume = 10 |issue = 12 |pages = 7829–7846 |bibcode = 2013BGeo...10.7829S }}</ref> Thus, macrofauna may alleviate N limitation by priming the remineralization of refractory N, reducing plants-microbe competition.<ref>{{cite journal |doi = 10.1111/j.1574-6941.2012.01400.x|title = Differential effects of microorganism-invertebrate interactions on benthic nitrogen cycling|year = 2012|last1 = Gilbertson|first1 = William W.|last2 = Solan|first2 = Martin|last3 = Prosser|first3 = James I.|journal = FEMS Microbiology Ecology|volume = 82|issue = 1|pages = 11–22|pmid = 22533682}}</ref><ref>{{cite journal |doi = 10.1042/BST0390315|title = Bioturbation: Impact on the marine nitrogen cycle|year = 2011|last1 = Laverock|first1 = Bonnie|last2 = Gilbert|first2 = Jack A.|last3 = Tait|first3 = Karen|last4 = Osborn|first4 = A. Mark|last5 = Widdicombe|first5 = Steve|journal = Biochemical Society Transactions|volume = 39|issue = 1|pages = 315–320|pmid = 21265795}}</ref> Such activity ultimately promotes N-recycling, plant assimilation and high N retention, as well as favours it loss by stimulating coupled nitrification and denitrification.<ref>{{cite journal |doi = 10.1002/lno.10724|title = Benthic N pathways in illuminated and bioturbated sediments studied with network analysis|year = 2018|last1 = Magri|first1 = M.|last2 = Benelli|first2 = S.|last3 = Bondavalli|first3 = C.|last4 = Bartoli|first4 = M.|last5 = Christian|first5 = R. R.|last6 = Bodini|first6 = A.|journal = Limnology and Oceanography|volume = 63|issue = S1|pages = S68–S84|bibcode = 2018LimOc..63S..68M}}</ref><ref name= Zilius2020 />

[[File:Nitrogen cycling in a mangrove fiddler crab holobiont.webp|thumb|upright=1.7|left|{{center|'''Nitrogen cycling in a mangrove fiddler crab holobiont'''{{hsp}}<ref name= Zilius2020>{{cite journal |doi = 10.1038/s41598-020-70834-0|title = N2 fixation dominates nitrogen cycling in a mangrove fiddler crab holobiont|year = 2020|last1 = Zilius|first1 = Mindaugas|last2 = Bonaglia|first2 = Stefano|last3 = Broman|first3 = Elias|last4 = Chiozzini|first4 = Vitor Gonsalez|last5 = Samuiloviene|first5 = Aurelija|last6 = Nascimento|first6 = Francisco J. A.|last7 = Cardini|first7 = Ulisse|last8 = Bartoli|first8 = Marco|journal = Scientific Reports|volume = 10|issue = 1|page = 13966|pmid = 32811860|pmc = 7435186}} [[File:CC-BY icon.svg|50px]] Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].</ref><br /><small>Dry weight of crab’s biofilm and mean dry weight<br />of incubated fiddler crab expressed as µmol N crab<sup>−1</sup> d<sup>−1</sup></small>}}]]

[[File:Thalassina anomala.JPG|thumb|upright=1.5|right|{{center| The [[scorpion mud lobster]] is found in some mangrove swamps. It lives in [[burrow]]s up to {{convert|2|m|ft|abbr=on}} deep, and is active at night. Its burrowing is important for the [[Biogeochemical cycle|recycling of nutrients]], bringing [[organic matter]] up from deep sediments.<ref name="Ng">{{cite book |title=A Guide to the Mangroves of Singapore |editor1=Peter K. L. Ng |editor2=N. Sivasothi |chapter=Mud lobster, ''Thalassina anomala'' |author=Kelvin K. P. Lim, Dennis H. Murphy, T. Morgany, N. Sivasothi, Peter K. L. Ng, B. C. Soong, Hugh T. W. Tan, K. S. Tan & T. K. Tan |year=1999 |isbn=981-04-1308-4 |publisher=[[Singapore Science Centre]] |chapter-url=http://mangrove.nus.edu.sg/guidebooks/text/2064.htm}}</ref><ref>{{cite web |url=http://www.naturia.per.sg/buloh/inverts/mudlobster.htm |title=Mud Lobster ''Thalassina anomala'' |author=Ria Tan |year=2001 |url-status=dead |archive-url=https://web.archive.org/web/20070827222018/http://www.naturia.per.sg/buloh/inverts/mudlobster.htm |archive-date=2007-08-27 }}</ref>]]

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Mangrove sediments are highly bioturbated by decapods such as crabs.<ref name=Kristensen2008>{{cite journal |doi = 10.1016/j.seares.2007.05.004|title = Mangrove crabs as ecosystem engineers; with emphasis on sediment processes|year = 2008|last1 = Kristensen|first1 = Erik|journal = Journal of Sea Research|volume = 59|issue = 1–2|pages = 30–43|bibcode = 2008JSR....59...30K}}</ref> Crab populations continuously rework sediment by constructing burrows, creating new niches, transporting or selectively grazing on sediment microbial communities.<ref name=Kristensen2008 /><ref>{{cite journal |doi = 10.1038/s41598-019-40315-0|title = Fiddler crab bioturbation determines consistent changes in bacterial communities across contrasting environmental conditions|year = 2019|last1 = Booth|first1 = Jenny Marie|last2 = Fusi|first2 = Marco|last3 = Marasco|first3 = Ramona|last4 = Mbobo|first4 = Tumeka|last5 = Daffonchio|first5 = Daniele|journal = Scientific Reports|volume = 9|issue = 1|page = 3749|pmid = 30842580|pmc = 6403291|bibcode = 2019NatSR...9.3749B}}</ref><ref name=Booth2019>{{cite journal |doi = 10.1038/s41396-017-0014-8|title = Multiple colonist pools shape fiddler crab-associated bacterial communities|year = 2018|last1 = Cuellar-Gempeler|first1 = Catalina|last2 = Leibold|first2 = Mathew A.|journal = The ISME Journal|volume = 12|issue = 3|pages = 825–837|pmid = 29362507|pmc = 5864236}}</ref><ref>{{cite journal |doi = 10.1016/j.jembe.2004.06.003|title = Impact of fiddler crab foraging and tidal inundation on an intertidal sandflat: Season-dependent effects in one tidal cycle|year = 2004|last1 = Reinsel|first1 = K.A.|journal = Journal of Experimental Marine Biology and Ecology|volume = 313|pages = 1–17}}</ref> In addition, crabs can affect organic matter turnover by assimilating leaves and producing finely fragmented faeces, or by carrying them into their burrows.<ref>{{cite journal |doi = 10.1016/j.jembe.2009.04.002|title = Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil|year = 2009|last1 = Nordhaus|first1 = Inga|last2 = Diele|first2 = Karen|last3 = Wolff|first3 = Matthias|journal = Journal of Experimental Marine Biology and Ecology|volume = 374|issue = 2|pages = 104–112}}</ref><ref>{{cite journal |doi = 10.1007/s00227-006-0597-5|title = Feeding ecology of the mangrove crab Ucides cordatus (Ocypodidae): Food choice, food quality and assimilation efficiency|year = 2007|last1 = Nordhaus|first1 = Inga|last2 = Wolff|first2 = Matthias|journal = Marine Biology|volume = 151|issue = 5|pages = 1665–1681|s2cid = 88582703}}</ref> Therefore, crabs are considered important ecosystem engineers shaping biogeochemical processes in intertidal muddy banks of mangroves.<ref>{{cite journal |doi = 10.1016/j.ecss.2011.03.002|title = Impact of crab bioturbation on benthic flux and nitrogen dynamics of Southwest Atlantic intertidal marshes and mudflats|year = 2011|last1 = Fanjul|first1 = Eugenia|last2 = Bazterrica|first2 = María C.|last3 = Escapa|first3 = Mauricio|last4 = Grela|first4 = María A.|last5 = Iribarne|first5 = Oscar|journal = Estuarine, Coastal and Shelf Science|volume = 92|issue = 4|pages = 629–638|bibcode = 2011ECSS...92..629F}}</ref><ref>{{cite journal |doi = 10.1038/srep16122|title = Carbon mineralization pathways and bioturbation in coastal Brazilian sediments|year = 2015|last1 = Quintana|first1 = Cintia O.|last2 = Shimabukuro|first2 = Maurício|last3 = Pereira|first3 = Camila O.|last4 = Alves|first4 = Betina G. R.|last5 = Moraes|first5 = Paula C.|last6 = Valdemarsen|first6 = Thomas|last7 = Kristensen|first7 = Erik|last8 = Sumida|first8 = Paulo Y. G.|journal = Scientific Reports|volume = 5|page = 16122|pmid = 26525137|pmc = 4630785|bibcode = 2015NatSR...516122Q}}</ref> In contrast to burrowing polychaetes or amphipods, the abundant Ocipodid crabs, mainly represented by fiddler crabs, do not permanently ventilate their burrows. These crabs may temporarily leave their burrows for surface activities18, or otherwise plug their burrow entrance during tidal inundation in order to trap air.<ref>{{cite journal |doi = 10.1016/0031-9384(94)90079-5|title = Burrow plugging in the crab Uca uruguayensis and its synchronization with photoperiod and tides|year = 1994|last1 = de la Iglesia|first1 = Horacio O.|last2 = Rodríguez|first2 = Enrique M.|last3 = Dezi|first3 = Rubén E.|journal = Physiology & Behavior|volume = 55|issue = 5|pages = 913–919|pmid = 8022913|s2cid = 7366251}}</ref> A recent study by Cuellar-Gempeler and Leibold17 showed that these crabs can be associated with a diverse microbial community, either on their carapace or in their gut.<ref name= Zilius2020 />

The exoskeleton of living animals, such as shells or carapaces, offers a habitat for microbial biofilms which are actively involved in different N-cycling pathways such as nitrification, denitrification and dissimilatory nitrate reduction to ammonium (DNRA).<ref>{{cite journal |doi = 10.1371/journal.pone.0185071|doi-access = free|title = Denitrification potential of the eastern oyster microbiome using a 16S rRNA gene based metabolic inference approach|year = 2017|last1 = Arfken|first1 = Ann|last2 = Song|first2 = Bongkeun|last3 = Bowman|first3 = Jeff S.|last4 = Piehler|first4 = Michael|journal = PLOS ONE|volume = 12|issue = 9|pages = e0185071|pmid = 28934286|pmc = 5608302|bibcode = 2017PLoSO..1285071A}}</ref><ref>{{cite journal |doi = 10.1016/j.marpolbul.2016.08.038|title = Living oysters and their shells as sites of nitrification and denitrification|year = 2016|last1 = Caffrey|first1 = Jane M.|last2 = Hollibaugh|first2 = James T.|last3 = Mortazavi|first3 = Behzad|journal = Marine Pollution Bulletin|volume = 112|issue = 1–2|pages = 86–90|pmid = 27567196}}</ref><ref>{{cite journal |doi = 10.1002/lno.10149|title = Copepod carcasses as microbial hot spots for pelagic denitrification|year = 2015|last1 = Glud|first1 = Ronnie N.|last2 = Grossart|first2 = Hans‐Peter|last3 = Larsen|first3 = Morten|last4 = Tang|first4 = Kam W.|last5 = Arendt|first5 = Kristine E.|last6 = Rysgaard|first6 = Søren|last7 = Thamdrup|first7 = Bo|last8 = Gissel Nielsen|first8 = Torkel|journal = Limnology and Oceanography|volume = 60|issue = 6|pages = 2026–2036|bibcode = 2015LimOc..60.2026G}}</ref><ref>{{cite journal |doi = 10.1111/j.1462-2920.2012.02823.x|title = Shell biofilm-associated nitrous oxide production in marine molluscs: Processes, precursors and relative importance|year = 2013|last1 = Heisterkamp|first1 = Ines M.|last2 = Schramm|first2 = Andreas|last3 = Larsen|first3 = Lone H.|last4 = Svenningsen|first4 = Nanna B.|last5 = Lavik|first5 = Gaute|last6 = De Beer|first6 = Dirk|last7 = Stief|first7 = Peter|journal = Environmental Microbiology|volume = 15|issue = 7|pages = 1943–1955|pmid = 22830624}}</ref><ref>{{cite journal |doi = 10.3354/meps13007|title = Nitrogen and phosphorus cycling in the digestive system and shell biofilm of the eastern oyster Crassostrea virginica|year = 2019|last1 = Ray|first1 = NE|last2 = Henning|first2 = MC|last3 = Fulweiler|first3 = RW|journal = Marine Ecology Progress Series|volume = 621|pages = 95–105|bibcode = 2019MEPS..621...95R|s2cid = 198261071}}</ref><ref>{{cite journal |doi = 10.1093/femsec/fiy144|title = Freshwater copepod carcasses as pelagic microsites of dissimilatory nitrate reduction to ammonium|year = 2018|last1 = Stief|first1 = Peter|last2 = Lundgaard|first2 = Ann Sofie Birch|last3 = Treusch|first3 = Alexander H.|last4 = Thamdrup|first4 = Bo|last5 = Grossart|first5 = Hans-Peter|last6 = Glud|first6 = Ronnie N.|journal = FEMS Microbiology Ecology|volume = 94|issue = 10|pmid = 30060193|pmc = 6084575}}</ref> Colonizing the carapace of crabs may be advantageous for specific bacteria, because of host activities such as respiration, excretion, feeding and horizontal and vertical migrations.<ref>{{cite journal |doi = 10.3389/fmicb.2012.00292|doi-access = free|title = The Second Skin: Ecological Role of Epibiotic Biofilms on Marine Organisms|year = 2012|last1 = Wahl|first1 = Martin|last2 = Goecke|first2 = Franz|last3 = Labes|first3 = Antje|last4 = Dobretsov|first4 = Sergey|last5 = Weinberger|first5 = Florian|journal = Frontiers in Microbiology|volume = 3|page = 292|pmid = 22936927|pmc = 3425911}}</ref> However, the ecological interactions between fiddler crabs and bacteria, their regulation and significance as well as their implications at scales spanning from the single individual to the ecosystem are not well understood.<ref name=Booth2019 /><ref>{{cite journal |doi = 10.1371/journal.pone.0130116|doi-access = free|title = The Link between Microbial Diversity and Nitrogen Cycling in Marine Sediments is Modulated by Macrofaunal Bioturbation|year = 2015|last1 = Yazdani Foshtomi|first1 = Maryam|last2 = Braeckman|first2 = Ulrike|last3 = Derycke|first3 = Sofie|last4 = Sapp|first4 = Melanie|last5 = Van Gansbeke|first5 = Dirk|last6 = Sabbe|first6 = Koen|last7 = Willems|first7 = Anne|last8 = Vincx|first8 = Magda|last9 = Vanaverbeke|first9 = Jan|journal = PLOS ONE|volume = 10|issue = 6|pages = e0130116|pmid = 26102286|pmc = 4477903|bibcode = 2015PLoSO..1030116Y}}</ref><ref name= Zilius2020 />

<gallery mode=packed style=float:left heights=180px caption="mangrove crabs">
File:Soldier crabs marching.jpg| Artist impression of small [[Mictyris longicarpus|blue soldier crabs]] marching across a mangrove flat{{hsp}}<ref>[https://australian.museum/learn/animals/crustaceans/soldier-crab/ Soldier Crab] ''Australian Museum''. Updated: 15 December 2020.</ref>
File:Mangrove crab.jpg|[[Mangrove crab]], possibly ''Neosarmatium meinerti''
File:Mangrove tree crab, Aratus pisonii.jpg|Mangrove tree crab, ''[[Aratus pisonii]]''
</gallery>
</gallery>



Revision as of 00:03, 7 November 2021

Mangrove ecosystem in the coastal intertidal zone [1]
Seagrass and oyster beds can inhabit the shallow subtidal zone

Mangrove forests are productive wetlands that occur in the intertidal zone along tropical and subtropical coasts.[2][3] Mangrove forests have attracted much research interest because of the various ecological functions of the mangrove ecosystems, including runoff and flood prevention, storage and recycling of nutrients and wastes, cultivation and energy conversion.[4] The forests live at the interface between the land, the ocean, and the atmosphere, and are centres for the flow of energy and matter between these systems. They are major blue carbon systems, storing considerable amounts of carbon in marine sediments, and thus becoming important regulators of climate change.[5][6]

Overview

The intertidal existence to which these trees are adapted represents the major limitation to the number of species able to thrive in their habitat. High tide brings in salt water, and when the tide recedes, solar evaporation of the seawater in the soil leads to further increases in salinity. The return of tide can flush out these soils, bringing them back to salinity levels comparable to that of seawater.[7][8]

At low tide, organisms are also exposed to increases in temperature and reduced moisture before being then cooled and flooded by the tide. Thus, for a plant to survive in this environment, it must tolerate broad ranges of salinity, temperature, and moisture, as well as several other key environmental factors—thus only a select few species make up the mangrove tree community.[8][7]

About 110 species are considered mangroves, in the sense of being trees that grow in such a saline swamp,[9] though only a few are from the mangrove plant genus, Rhizophora. However, a given mangrove swamp typically features only a small number of tree species. It is not uncommon for a mangrove forest in the Caribbean to feature only three or four tree species. For comparison, the tropical rainforest biome contains thousands of tree species, but this is not to say mangrove forests lack diversity. Though the trees themselves are few in species, the ecosystem that these trees create provides a home (habitat) for a great variety of other species, including as many as 174 species of marine megafauna.[10]

Mangrove plants require a number of physiological adaptations to overcome the problems of low environmental oxygen levels, high salinity, and frequent tidal flooding. Each species has its own solutions to these problems; this may be the primary reason why, on some shorelines, mangrove tree species show distinct zonation. Small environmental variations within a mangal may lead to greatly differing methods for coping with the environment. Therefore, the mix of species is partly determined by the tolerances of individual species to physical conditions, such as tidal flooding and salinity, but may also be influenced by other factors, such as crabs preying on plant seedlings.[11]

Once established, mangrove roots provide an oyster habitat and slow water flow, thereby enhancing sediment deposition in areas where it is already occurring. The fine, anoxic sediments under mangroves act as sinks for a variety of heavy (trace) metals which colloidal particles in the sediments have concentrated from the water. Mangrove removal disturbs these underlying sediments, often creating problems of trace metal contamination of seawater and organisms of the area.[12]

Mangrove swamps protect coastal areas from erosion, storm surge (especially during tropical cyclones), and tsunamis.[13][14][15] They limit high-energy wave erosion mainly during events such as storm surges and tsunamis.[16] The mangroves' massive root systems are efficient at dissipating wave energy.[17] Likewise, they slow down tidal water enough so that its sediment is deposited as the tide comes in, leaving all except fine particles when the tide ebbs.[18] In this way, mangroves build their environments.[13] Because of the uniqueness of mangrove ecosystems and the protection against erosion they provide, they are often the object of conservation programs,[8] including national biodiversity action plans.[14]

Distribution

Global distribution of mangrove forests
Distribution of mangroves and carbonate sediments around South-East Asia and the South Pacific
Distribution of deltaic, estuarine, lagoonal and open coast mangrove types, and approximate extent of carbonate sedimentary settings in (ai) the South Asia, (aii) Southeast Asia and (aiii) East Asia regions and (bi) the Australia and New Zealand and (bii) Pacific Islands regions. Bar charts represent the percentage change in area of the different types between 1996 and 2016 at the regional scale. *Value truncated for display, actual value − 33.2%.[19][20]}}

The largest mangrove forest in the world is in the Sundarbans. The Sundarban forest lies in the vast delta on the Bay of Bengal formed by the super confluence of the Hooghly, Padma (both are distributaries of Ganges), Brahmaputra and Meghna rivers across southern Bangladesh. The seasonally flooded Sundarbans freshwater swamp forests lie inland from the mangrove forests on the coastal fringe. The forest covers 10,000 km2 (3,900 sq mi) of which about 6,000 km2 (2,300 sq mi) are in Bangladesh. The Indian part of Sundarbans is estimated to be about 4,110 km2 (1,590 sq mi), of which about 1,700 km2 (660 sq mi) is occupied by water bodies in the forms of river, canals and creeks of width varying from a few metres to several kilometres.[citation needed]

The Sundarbans is intersected by a complex network of tidal waterways, mudflats and small islands of salt-tolerant mangrove forests. The interconnected network of waterways makes almost every corner of the forest accessible by boat. The area is known for the Bengal tiger (Panthera tigris), as well as numerous fauna including species of birds, spotted deer, crocodiles and snakes. The fertile soils of the delta have been subject to intensive human use for centuries, and the ecoregion has been mostly converted to intensive agriculture, with few enclaves of forest remaining. The remaining forests, taken together with the Sundarbans mangroves, are important habitat for the endangered tiger. Additionally, the Sundarbans serves a crucial function as a protective barrier for the millions of inhabitants in and around Khulna and Mongla against the floods that result from the cyclones.[citation needed]

Sundarbans is located in South Asia
Sundarbans
Sundarbans
Sundarbans (South Asia)

The forest ecosystem

The unique ecosystem found in the intricate mesh of mangrove roots offers a quiet marine habitat for young organisms.[21] In areas where roots are permanently submerged, the organisms they host include algae, barnacles, oysters, sponges, and bryozoans, which all require a hard surface for anchoring while they filter-feed. Shrimps and mud lobsters use the muddy bottoms as their home.[22] Mangrove crabs eat the mangrove leaves, adding nutrients to the mangal mud for other bottom feeders.[23] In at least some cases, the export of carbon fixed in mangroves is important in coastal food webs.[24]

Mangrove plantations in Vietnam, Thailand, Philippines, and India host several commercially important species of fish and crustaceans.[25]

Mangrove forests can decay into peat deposits because of fungal and bacterial processes as well as by the action of termites. It becomes peat in good geochemical, sedimentary, and tectonic conditions.[26] The nature of these deposits depends on the environment and the types of mangroves involved. In Puerto Rico, the red, white, and black mangroves occupy different ecological niches and have slightly different chemical compositions, so the carbon content varies between the species, as well between the different tissues of the plant (e.g., leaf matter versus roots).[26]

In Puerto Rico, there is a clear succession of these three trees from the lower elevations, which are dominated by red mangroves, to farther inland with a higher concentration of white mangroves.[26] Mangrove forests are an important part of the cycling and storage of carbon in tropical coastal ecosystems.[26] Knowing this, scientists seek to reconstruct the environment and investigate changes to the coastal ecosystem over thousands of years using sediment cores.[27] However, an additional complication is the imported marine organic matter that also gets deposited in the sediment due to the tidal flushing of mangrove forests.[26]

Termites play an important role in the formation of peat from mangrove materials.[26] They process fallen leaf litter, root systems and wood from mangroves into peat to build their nests.[26] Termites stabilise the chemistry of this peat and represent approximately 2% of above ground carbon storage in mangroves.[26] As the nests are buried over time this carbon is stored in the sediment and the carbon cycle continues.[26]

Mangroves are an important source of blue carbon. Globally, mangroves stored 4.19 Gt (9.2×1012 lb) of carbon in 2012.[28] Two percent of global mangrove carbon was lost between 2000 and 2012, equivalent to a maximum potential of 0.316996250 Gt (6.9885710×1011 lb) of CO2 emissions.[28]

Globally, mangroves have been shown to provide measurable economic protections to coastal communities affected by tropical storms.[29]

Mangrove birds

Heterogeneity in landscape ecology is a measure of how different parts of a landscape are from one another. It can manifest in an ecosystem from the abiotic or biotic characteristics of the environment. For example, coastal mangrove forests are located at the land-sea interface, so their functioning is influenced by abiotic factors such as tides, as well as biotic factors such as the extent and configuration of adjacent vegetation.[30] For forest birds, tidal inundation means that the availability of many mangrove resources fluctuates daily, suggesting foraging flexibility is likely to be important. Mangroves also offer estuarine prey items, such as mudskippers and crabs. that are not found in terrestrial forest types. Further, mangroves are often situated in a complex mosaic of adjacent vegetation types such as grasslands, saltmarshes, and woodlands, and this can mean that flexibility in foraging strategy and choice of foraging habitat may be advantageous for highly mobile forest birds.[30] Relative to other forest types, mangroves support few bird species that are obligate habitat (mangrove) specialists and instead host many species with generalised foraging niches.[31][32][33][30]

Mangrove crabs

Mangrove forests are among the most productive and diverse ecosystems on the planet, despite limited nitrogen (N) availability. Under such conditions, animal-microbe associations (holobionts) are often key to ecosystem functioning. Here, we investigated the role of fiddler crabs and their carapace-associated microbial biofilm as hotspots of microbial N transformations and sources of N within the mangrove ecosystem.[34]

Among coastal ecosystems, mangrove forests are of great importance as they account for three quarters of the tropical coastline and provide different ecosystem services.[35][36] Mangrove ecosystems generally act as a net sink of carbon, although they release organic matter to the sea in the form of dissolved refractory macromolecules, leaves, branches and other debris.[37][38] In pristine environments, mangroves are among the most productive ecosystems on the planet, despite growing in tropical waters that are often nutrient depleted.[39] The refractory nature of the organic matter produced and retained in mangroves can slow the recycling of nutrients, particularly of nitrogen (N).[37][40] Nitrogen limitation in such systems may be overcome by microbial dinitrogen (N2) fixation when combined with high rates of bioturbation by macrofauna.[41][42][34]

Bioturbation by macrofauna affect N availability and multiple N-related microbial processes through sediment reworking, burrow construction and bioirrigation, feeding and excretion.[43] Macrofauna mix old and fresh organic matter, extend oxic–anoxic sediment interfaces, increase the availability of energy-yielding electron acceptors and increase N turnover via direct excretion.[44][45] Thus, macrofauna may alleviate N limitation by priming the remineralization of refractory N, reducing plants-microbe competition.[46][47] Such activity ultimately promotes N-recycling, plant assimilation and high N retention, as well as favours it loss by stimulating coupled nitrification and denitrification.[48][34]

Nitrogen cycling in a mangrove fiddler crab holobiont[34]
Dry weight of crab’s biofilm and mean dry weight
of incubated fiddler crab expressed as µmol N crab−1 d−1
The scorpion mud lobster is found in some mangrove swamps. It lives in burrows up to 2 m (6.6 ft) deep, and is active at night. Its burrowing is important for the recycling of nutrients, bringing organic matter up from deep sediments.[49][50]

Mangrove sediments are highly bioturbated by decapods such as crabs.[51] Crab populations continuously rework sediment by constructing burrows, creating new niches, transporting or selectively grazing on sediment microbial communities.[51][52][53][54] In addition, crabs can affect organic matter turnover by assimilating leaves and producing finely fragmented faeces, or by carrying them into their burrows.[55][56] Therefore, crabs are considered important ecosystem engineers shaping biogeochemical processes in intertidal muddy banks of mangroves.[57][58] In contrast to burrowing polychaetes or amphipods, the abundant Ocipodid crabs, mainly represented by fiddler crabs, do not permanently ventilate their burrows. These crabs may temporarily leave their burrows for surface activities18, or otherwise plug their burrow entrance during tidal inundation in order to trap air.[59] A recent study by Cuellar-Gempeler and Leibold17 showed that these crabs can be associated with a diverse microbial community, either on their carapace or in their gut.[34]

The exoskeleton of living animals, such as shells or carapaces, offers a habitat for microbial biofilms which are actively involved in different N-cycling pathways such as nitrification, denitrification and dissimilatory nitrate reduction to ammonium (DNRA).[60][61][62][63][64][65] Colonizing the carapace of crabs may be advantageous for specific bacteria, because of host activities such as respiration, excretion, feeding and horizontal and vertical migrations.[66] However, the ecological interactions between fiddler crabs and bacteria, their regulation and significance as well as their implications at scales spanning from the single individual to the ecosystem are not well understood.[53][67][34]

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