Mangrove

{{Short description|Shrub growing in brackish water}}

{{Other uses}}

{{Use dmy dates|date=July 2022}}

File:Sonneratia alba - Manado (2).JPGs along marine coasts.]]

A mangrove is a shrub or tree that grows mainly in coastal saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen and remove salt, allowing them to tolerate conditions that kill most plants. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse due to convergent evolution in several plant families. They occur worldwide in the tropics and subtropics and even some temperate coastal areas, mainly between latitudes 30° N and 30° S, with the greatest mangrove area within 5° of the equator. Mangrove plant families first appeared during the Late Cretaceous to Paleocene epochs and became widely distributed in part due to the movement of tectonic plates. The oldest known fossils of mangrove palm date to 75 million years ago.

Mangroves are salt-tolerant (halophytic) and are adapted to live in harsh coastal conditions. They contain a complex salt filtration system and a complex root system to cope with saltwater immersion and wave action. They are adapted to the low-oxygen conditions of waterlogged mud,{{cite journal |last1=Flowers |first1=T. J. |last2=Colmer |first2=T. D. |year=2015 |title=Plant salt tolerance: adaptations in halophytes |journal=Annals of Botany |volume=115 |issue=3 |pages=327–331 |doi=10.1093/aob/mcu267 |pmid=25844430 |pmc=4332615 |url=}} but are most likely to thrive in the upper half of the intertidal zone.{{cite journal |last1=Zimmer |first1=Katarina |title=Many mangrove restorations fail. Is there a better way? |journal=Knowable Magazine |date=22 July 2021 |doi=10.1146/knowable-072221-1 |url=https://knowablemagazine.org/article/food-environment/2021/many-mangrove-restorations-fail |access-date=11 August 2021 |doi-access=free}}

The mangrove biome, often called the mangrove forest or mangal, is a distinct saline woodland or shrubland habitat characterized by depositional coastal environments, where fine sediments (often with high organic content) collect in areas protected from high-energy wave action. Mangrove forests serve as vital habitats for a diverse array of aquatic species, offering a unique ecosystem that supports the intricate interplay of marine life and terrestrial vegetation. The saline conditions tolerated by various mangrove species range from brackish water, through pure seawater (3 to 4% salinity), to water concentrated by evaporation to over twice the salinity of ocean seawater (up to 9% salinity).{{cite web|url=http://www.nhmi.org/mangroves/phy.htm |title=Morphological and Physiological Adaptations: Florida mangrove website |publisher=Nhmi.org |access-date=8 February 2012 |url-status=dead |archive-url=https://web.archive.org/web/20120204222702/http://www.nhmi.org/mangroves/phy.htm |archive-date=4 February 2012 }}{{cite book |last1=Primavera |first1=J. H. |last2=Savaris |first2=J. P. |last3=Bajoyo |first3=B. E. |last4=Coching |first4=J. D. |last5=Curnick |first5=D. J. |last6=Golbeque |first6=R. L. |last7=Guzman |first7=A. T. |last8=Henderin |first8=J. Q. |last9=Joven |first9=R. V. |last10=Loma |first10=R. A. |last11=Koldewey |first11=H. J. |title=Manual on community-based mangrove rehabilitation |series=Mangrove Manual |date=2012 |publisher=The Zoological Society of London ZSL |url=https://www.zsl.org/sites/default/files/media/2014-05/Manual%20on%20Community-Based%20Mangrove%20Rehabilitation.pdf |access-date=15 August 2021 |archive-date=1 January 2016 |archive-url=https://web.archive.org/web/20160101053522/http://www.zsl.org/sites/default/files/media/2014-05/Manual%20on%20Community-Based%20Mangrove%20Rehabilitation.pdf |url-status=dead }}

Beginning in 2010, remote sensing technologies and global data have been used to assess areas, conditions and deforestation rates of mangroves around the world.{{cite journal |last1=Bunting |first1=P. |last2=Rosenqvist |first2=A. |last3=Lucas |first3=R. |last4=Rebelo |first4=L.-M. |last5=Hilarides |first5=L. |last6=Thomas |first6=N. |last7=Hardy |first7=A. |last8=Itoh |first8=T. |last9=Shimada |first9=M. |last10=Finlayson |first10=C. |title=The Global Mangrove Watch—A New 2010 Global Baseline of Mangrove Extent |journal=Remote Sensing |date=2018 |volume=10 |issue=10 |pages=1669 |doi=10.3390/rs10101669 |bibcode=2018RemS...10.1669B |doi-access=free}}{{cite journal |last1=Giri |first1=C. |last2=Ochieng |first2=E. |last3=Tieszen |first3=L. L. |last4=Zhu |first4=Z. |last5=Singh |first5=A. |last6=Loveland |first6=T. |last7=Masek |first7=J. |last8=Duke |first8=N. |title=Status and distribution of mangrove forests of the world using earth observation satellite data: Status and distributions of global mangroves |journal=Global Ecology and Biogeography |date=2011 |volume=20 |issue=1 |pages=154–159 |doi=10.1111/j.1466-8238.2010.00584.x |doi-access=free}}{{cite journal |last1=Friess |first1=D. A. |last2=Rogers |first2=K. |last3=Lovelock |first3=C. E. |last4=Krauss |first4=K. W. |last5=Hamilton |first5=S. E. |last6=Lee |first6=S. Y. |last7=Lucas |first7=R. |last8=Primavera |first8=J. |last9=Rajkaran |first9=A. |last10=Shi |first10=S. |title=The State of the World's Mangrove Forests: Past, Present, and Future |journal=Annual Review of Environment and Resources |date=2019 |volume=44 |issue=1 |pages=89–115 |doi=10.1146/annurev-environ-101718-033302 |doi-access=free}} In 2018, the Global Mangrove Watch Initiative released a new global baseline which estimates the total mangrove forest area of the world as of 2010 at {{cvt|137600|km2}}, spanning 118 countries and territories. A 2022 study on losses and gains of tidal wetlands estimates a {{cvt|3700|km2}} net decrease in global mangrove extent from 1999 to 2019.{{cite journal |last1=Murray |first1=N. J. |last2=Worthington |first2=T. A. |last3=Bunting |first3=P. |last4=Duce |first4=S. |last5=Hagger |first5=V. |last6=Lovelock |first6=C. E. |last7=Lucas |first7=R. |last8=Saunders |first8=M. I. |last9=Sheaves |first9=M. |last10=Spalding |first10=M. |last11=Waltham |first11=N. J. |last12=Lyons |first12=M. B. |title=High-resolution mapping of losses and gains of Earth's tidal wetlands |journal=Science |date=2022 |volume=376 |issue=6594 |pages=744–749 |doi=10.1126/science.abm9583|pmid=35549414 |bibcode=2022Sci...376..744M |s2cid=248749118 |url=https://www.repository.cam.ac.uk/handle/1810/337253|doi-access=free |hdl=2160/55fdc0d4-aa3e-433f-8a88-2098b1372ac5 |hdl-access=free }} Mangrove loss continues due to human activity, with a global annual deforestation rate estimated at 0.16%, and per-country rates as high as 0.70%. Degradation in quality of remaining mangroves is also an important concern.

There is interest in mangrove restoration for several reasons. Mangroves support sustainable coastal and marine ecosystems. They protect nearby areas from tsunamis and extreme weather events. Mangrove forests are also effective at carbon sequestration and storage.{{Cite web |title=New Satellite-Based maps of Mangrove heights | access-date = 15 May 2019 |date=2019 |url=https://earthobservatory.nasa.gov/images/144573/new-satellite-based-maps-of-mangrove-heights?_ga=2.242896961.239110103.1557888044-1148588132.1557888044 |first1=Carol |last1=R. |first2=M. |last2=Carlowicz}}{{cite journal|last1= Simard|first1=M.|last2=Fatoyinbo|first2= L. |last3=Smetanka |first3=C.|last4=Rivera-Monroy|first4=V. H. |last5=Castañeda-Moya|first5=E. |last6=Thomas |first6=N. |last7=Van der Stocken|first7=T.|title=Mangrove canopy height globally related to precipitation, temperature and cyclone frequency |journal= Nature Geoscience |volume=12|issue=1|year=2018|pages=40–45 |doi=10.1038/s41561-018-0279-1 |hdl=2060/20190029179|s2cid=134827807|hdl-access=free}} The success of mangrove restoration may depend heavily on engagement with local stakeholders, and on careful assessment to ensure that growing conditions will be suitable for the species chosen.

The International Day for the Conservation of the Mangrove Ecosystem is celebrated every year on 26 July.{{Cite web |title=International Day for the Conservation of the Mangrove Ecosystem |url=https://www.unesco.org/en/days/mangrove-ecosystem-conservation?TSPD_101_R0=080713870fab200040cc16c5f9135ee1e980e337118e23f8f0052aa91f72e343e722a66985cbe18008c5863437144800f8f8fe53986f7a675dffd9ccac6111d49e4b3179755fa00cd692e29021fad73a62b94bb5cdf481e97bc6540c36bb8cc2268bc4db3a44bd9fa03a3763ae2c830b1e7e6068a581610b |access-date=9 June 2023 |website=UNESCO}}

Etymology

File:Mangrove roots at low tide.jpg

File:Mangroves in Kannur, India.jpg

Etymology of the English term mangrove can only be speculative and is disputed.{{cite book |last1=Saenger |first1=P. |title=Mangrove ecology, silviculture, and conservation |date=2013 |publisher=Springer Science & Business Media |location= |isbn=9789401599627 |edition=Reprint of 2002 |url=https://books.google.com/books?id=WUbrCAAAQBAJ&pg=PA2}}{{rp|1–2}}

The term may have come to English from the Portuguese {{lang|pt|mangue}} or the Spanish

{{lang|es|mangle}}. Further back, it may be traced to South America and Cariban and Arawakan languages{{cite book |last1=Görlach |first1=M. |title=English Words Abroad |date=1 January 2003 |publisher=John Benjamins Publishing |page=59 |isbn=9027223319 |url=https://books.google.com/books?id=Ph0kR1eiEJQC&pg=PA59 |access-date=13 August 2021}} such as Taíno.{{cite book|author= Rafinesque, C. S.|author-link= Constantine Samuel Rafinesque|title=The American Nations|volume= 1|url= https://books.google.com/books?id=XYlHAAAAIAAJ&pg=PA244|year= 1836|publisher=C. S. Rafinesque|page=244}} Other possibilities include the Malay language {{lang|ms|manggi-manggi}}

The English usage may reflect a corruption via folk etymology of the words mangrow and grove.{{cite book |last1=Weekley |first1=Ernest |title=An Etymological Dictionary of Modern English |date=1967 |publisher=Dover |volume=2 |isbn=9780486122861 |edition=Reprint of 1921 |url=https://books.google.com/books?id=VigsBojU2c4C&pg=PA890 |access-date=13 August 2021}}

The word "mangrove" is used in at least three senses:

  • Most broadly to refer to the habitat and entire plant assemblage or mangal,{{cite journal |last1=Macnae |first1=W. |title=A General Account of the Fauna and Flora of Mangrove Swamps and Forests in the Indo-West-Pacific Region |journal=Advances in Marine Biology |date=1969 |volume=6 |pages=73–270 |doi=10.1016/S0065-2881(08)60438-1 |isbn=9780120261062 |url=https://books.google.com/books?id=UefCHv9EU5IC&pg=PA75 |access-date=13 August 2021}}{{cite book | last=Hogarth | first=Peter J. | title=The biology of mangroves and seagrasses | publisher=Oxford university press | publication-place=Oxford | date=2015 | isbn=978-0-19-871654-9}} for which the terms mangrove forest biome and mangrove swamp are also used;
  • To refer to all trees and large shrubs in a mangrove swamp; and
  • Narrowly to refer only to mangrove trees of the genus Rhizophora of the family Rhizophoraceae.{{Cite book|last=Austin|first=D. F. |title=Florida Ethnobotany |date=2004 |publisher=CRC Press |url=https://books.google.com/books?id=eS7lX_rC3GEC|isbn=978-0-203-49188-1}}

Biology

According to Hogarth (2015), among the recognized mangrove species there are about 70 species in 20 genera from 16 families that constitute the "true mangroves" – species that occur almost exclusively in mangrove habitats. Demonstrating convergent evolution, many of these species found similar solutions to the tropical conditions of variable salinity, tidal range (inundation), anaerobic soils, and intense sunlight. Plant biodiversity is generally low in a given mangrove. The greatest biodiversity of mangroves occurs in Southeast Asia, particularly in the Indonesian archipelago.{{cite web |url=http://maps.grida.no/go/graphic/distribution_of_coral_mangrove_and_seagrass_diversity/ |title=Distribution of coral, mangrove and seagrass diversity |publisher=Maps.grida.no |access-date=8 February 2012 |url-status=dead |archive-url=https://web.archive.org/web/20100305202831/http://maps.grida.no/go/graphic/distribution_of_coral_mangrove_and_seagrass_diversity |archive-date=5 March 2010 }}

File:Mangrove (cropped).jpg

=Adaptations to low oxygen=

The red mangrove (Rhizophora mangle) survives in the most inundated areas, props itself above the water level with stilt or prop roots and then absorbs air through lenticels in its bark.{{cite web |title=Red mangrove |url=https://www.daf.qld.gov.au/business-priorities/fisheries/habitats/marine-plants-including-mangroves/common-mangroves/red-mangrove |website=Department of Agriculture and Fisheries, Queensland Government |date=January 2013 |access-date=13 August 2021}}

The black mangrove (Avicennia germinans) lives on higher ground and develops many specialized root-like structures called pneumatophores, which stick up out of the soil like straws for breathing.{{cite web |title=Black Mangrove (Avicennia germinans) |url=https://environment.bm/black-mangrove |website=The Department of Environment and Natural Resources, Government of Bermuda |access-date=13 August 2021}}{{cite web |title=Morphological and Physiological Adaptations |url=https://www.nhmi.org/mangroves/phy.htm |website=Newfound Harbor Marine Institute |access-date=13 August 2021}}

These "breathing tubes" typically reach heights of up to {{Cvt|30|cm|in}}, and in some species, over {{Cvt|3|m}}. The roots also contain wide aerenchyma to facilitate transport within the plants.{{citation needed|date=June 2021}}

=Nutrient uptake=

Because the soil is perpetually waterlogged, little free oxygen is available. Anaerobic bacteria liberate nitrogen gas, soluble ferrum (iron), inorganic phosphates, sulfides, and methane, which make the soil much less nutritious.{{Citation needed|date=November 2011}} Pneumatophores (aerial roots) allow mangroves to absorb gases directly from the atmosphere, and other nutrients such as iron, from the inhospitable soil. Mangroves store gases directly inside the roots, processing them even when the roots are submerged during high tide.

File:Saltcrystals on avicennia marina var resinifera leaves.JPG leaf]]

=Limiting salt intake=

Red mangroves exclude salt by having significantly impermeable roots that are highly suberised (impregnated with suberin), acting as an ultrafiltration mechanism to exclude sodium salts from the rest of the plant.{{Citation needed|date=June 2023}} One study found that roots of the Indian mangrove Avicennia officinalis exclude 90% to 95% of the salt in water taken up by the plant, depositing the excluded salt in the cortex of the root. An increase in the production of suberin and in the activity of a gene regulating cytochrome P450 were observed in correlation with an increase in the salinity of the water to which the plant was exposed.{{Cite journal |last1=Krishnamurthy |first1=Pannaga |last2=Jyothi-Prakash |first2=Pavithra A. |last3=Qin |first3=Lin |last4=He |first4=Jie |last5=Lin |first5=Qingsong |last6=Loh |first6=Chiang-Shiong |last7=Kumar |first7=Prakash P. |date=July 2014 |title=Role of root hydrophobic barriers in salt exclusion of a mangrove plant Avicennia officinalis |journal=Plant, Cell & Environment |language=en |volume=37 |issue=7 |pages=1656–1671 |doi=10.1111/pce.12272|pmid=24417377 |doi-access=free }} In a frequently cited concept that has become known as the "sacrificial leaf", salt which does accumulate in the shoot (sprout) then concentrates in old leaves, which the plant then sheds. However, recent research on the Red mangrove Rhizophora mangle suggests that the older, yellowing leaves have no more measurable salt content than the other, greener leaves.{{cite web|last=Gray|first=L. Joseph |year=2010 |title=Sacrificial leaf hypothesis of mangroves |url=http://www.glomis.com/ej/pdf/EJ_8-4.pdf |work=ISME/GLOMIS Electronic Journal |publisher=GLOMIS |access-date=21 January 2012 |display-authors=etal}}

File:Pneumatophore overkill - grey mangrove.JPG|Pneumatophorous aerial roots of the grey mangrove (Avicennia marina)

File:Plody mangrovnika (Rhizophora mangle).jpg|Vivipary in Rhizophora mangle seeds

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=Limiting water loss=

File:Water filtration in mangrove roots.webp. (a) Schematic of the root. The outermost layer is composed of three layers. The root is immersed in NaCl solution. (b) Water passes through the outermost layer when a negative suction pressure is applied across the outermost layer. The Donnan potential effect repels Cl ions from the first sublayer of the outermost layer. Na+ ions attach to the first layer to satisfy the electro-neutrality requirement and salt retention eventually occurs.{{cite journal | last1=Kim | first1=Kiwoong | last2=Seo | first2=Eunseok | last3=Chang | first3=Suk-Kyu | last4=Park | first4=Tae Jung | last5=Lee | first5=Sang Joon | title=Novel water filtration of saline water in the outermost layer of mangrove roots | journal=Scientific Reports | publisher=Springer Science and Business Media LLC | volume=6 | issue=1 | date=5 February 2016 | page=20426 | issn=2045-2322 | doi=10.1038/srep20426| pmid=26846878 | pmc=4742776 | bibcode=2016NatSR...620426K }} 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].]]

Because of the limited fresh water available in salty intertidal soils, mangroves limit the amount of water they lose through their leaves. They can restrict the opening of their stomata (pores on the leaf surfaces, which exchange carbon dioxide gas and water vapor during photosynthesis). They also vary the orientation of their leaves to avoid the harsh midday sun and so reduce evaporation from the leaves. A captive red mangrove grows only if its leaves are misted with fresh water several times a week, simulating frequent tropical rainstorms.{{cite web |author=Calfo |first=Anthony |year=2006 |title=Mangroves for the Marine Aquarium |url=http://www.reefkeeping.com/issues/2004-12/ac/feature/index.php |url-status=live |archive-url=https://web.archive.org/web/20220201145128/http://reefkeeping.com/issues/2004-12/ac/feature/index.php |archive-date=1 February 2022 |access-date=8 February 2012 |website=Reefkeeping |publisher=Reef Central}}

=Filtration of seawater=

A 2016 study by Kim et al. investigated the biophysical characteristics of sea water filtration in the roots of the mangrove Rhizophora stylosa from a plant hydrodynamic point of view. R. stylosa can grow even in saline water and the salt level in its roots is regulated within a certain threshold value through filtration. The root possesses a hierarchical, triple layered pore structure in the epidermis and most Na+ ions are filtered at the first sublayer of the outermost layer. The high blockage of Na+ ions is attributed to the high surface zeta potential of the first layer. The second layer, which is composed of macroporous structures, also facilitates Na+ ion filtration. The study provides insights into the mechanism underlying water filtration through halophyte roots and could serve as a basis for the development of a bio-inspired method of desalination.

Uptake of Na+ ions is desirable for halophytes to build up osmotic potential, absorb water and sustain turgor pressure. However, excess Na+ ions may work on toxic element. Therefore, halophytes try to adjust salinity delicately between growth and survival strategies. In this point of view, a novel sustainable desalination method can be derived from halophytes, which are in contact with saline water through their roots. Halophytes exclude salt through their roots, secrete the accumulated salt through their aerial parts and sequester salt in senescent leaves and/or the bark.Tomlinson, P. The botany of mangroves. [116–130] (Cambridge University Press, Cambridge, 1986).{{cite journal |doi = 10.1016/S0022-0981(98)00131-2|title = Dynamics of element contents during the development of hypocotyles and leaves of certain mangrove species|year = 1999|last1 = Zheng|first1 = Wen-Jiao|last2 = Wang|first2 = Wen-Qing|last3 = Lin|first3 = Peng|journal = Journal of Experimental Marine Biology and Ecology|volume = 233|issue = 2|pages = 247–257| bibcode=1999JEMBE.233..247Z }}{{cite journal |doi = 10.1007/s00468-010-0417-x|title = Salt tolerance mechanisms in mangroves: A review|year = 2010|last1 = Parida|first1 = Asish Kumar|last2 = Jha|first2 = Bhavanath|journal = Trees|volume = 24|issue = 2|pages = 199–217| bibcode=2010Trees..24..199P |s2cid = 3036770}} Mangroves are facultative halophytes and Bruguiera is known for its special ultrafiltration system that can filter approximately 90% of Na+ions from the surrounding seawater through the roots.{{cite journal |doi = 10.1111/j.1399-3054.1968.tb07248.x|title = How Mangroves Desalinate Seawater|year = 1968|last1 = Scholander|first1 = P. F.|journal = Physiologia Plantarum|volume = 21|pages = 251–261}}{{cite journal |doi = 10.1104/pp.41.3.529|title = Sap Concentrations in Halophytes and Some Other Plants|year = 1966|last1 = Scholander|first1 = P. F.|last2 = Bradstreet|first2 = Edda D.|last3 = Hammel|first3 = H. T.|last4 = Hemmingsen|first4 = E. A.|journal = Plant Physiology|volume = 41|issue = 3|pages = 529–532|pmid = 5906381|pmc = 1086377}} The species also exhibits a high rate of salt rejection. The water-filtering process in mangrove roots has received considerable attention for several decades.{{cite journal |doi = 10.1111/j.1469-8137.1982.tb03338.x|title = Physiology of Salt Excretion in the Mangrove Avicennia Marina (Forsk.) Vierh|year = 1982|last1 = Drennan|first1 = Philippa|last2 = Pammenter|first2 = N. W.|journal = New Phytologist|volume = 91|issue = 4|pages = 597–606|doi-access = free}}{{cite journal |doi = 10.1016/S0367-2530(17)30013-0|title = Effect of high external Na Cl concentration on the osmolality of xylem sap, leaf tissue and leaf glands secretion of the mangrove Avicennia germinans (L.) L|year = 2001|last1 = Sobrado|first1 = M.A.|journal = Flora|volume = 196| issue=1 |pages = 63–70| bibcode=2001FMDFE.196...63S }} Morphological structures of plants and their functions have been evolved through a long history to survive against harsh environmental conditions.{{cite journal |doi = 10.1016/j.pbi.2006.05.014|title = Crosstalk between abiotic and biotic stress responses: A current view from the points of convergence in the stress signaling networks|year = 2006|last1 = Fujita|first1 = Miki|last2 = Fujita|first2 = Yasunari|last3 = Noutoshi|first3 = Yoshiteru|last4 = Takahashi|first4 = Fuminori|last5 = Narusaka|first5 = Yoshihiro|last6 = Yamaguchi-Shinozaki|first6 = Kazuko|last7 = Shinozaki|first7 = Kazuo|journal = Current Opinion in Plant Biology|volume = 9|issue = 4|pages = 436–442|pmid = 16759898| bibcode=2006COPB....9..436F | s2cid=31166870 }}

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=Increasing survival of offspring=

File:One week old Mangarove, Qatif, Saudi Arabia, Late August 2020 2 (cropped).jpg

{{Unreferenced section|date=October 2021}}

In this harsh environment, mangroves have evolved a special mechanism to help their offspring survive. Mangrove seeds are buoyant and are therefore suited to water dispersal. Unlike most plants, whose seeds germinate in soil, many mangroves (e.g. red mangrove) are viviparous,{{Citation |last=Hogarth |first=P. J. |title=Mangrove Ecosystems☆ |date=2017-01-01 |work=Reference Module in Life Sciences |url=https://www.sciencedirect.com/science/article/pii/B9780128096338022093 |access-date=2024-03-01 |publisher=Elsevier |doi=10.1016/b978-0-12-809633-8.02209-3 |isbn=978-0-12-809633-8}} meaning their seeds germinate while still attached to the parent tree. Once germinated, the seedling grows either within the fruit (e.g. Aegialitis, Avicennia and Aegiceras), or out through the fruit (e.g. Rhizophora, Ceriops, Bruguiera and Nypa) to form a propagule (a ready-to-go seedling) which can produce its own food via photosynthesis.

The mature propagule then drops into the water, which can transport it great distances. Propagules can survive desiccation and remain dormant for over a year before arriving in a suitable environment. Once a propagule is ready to root, its density changes so that the elongated shape now floats vertically rather than horizontally. In this position, it is more likely to lodge in the mud and root. If it does not root, it can alter its density and drift again in search of more favorable conditions.

Taxonomy and evolution

The following listings, based on Tomlinson, 2016, give the mangrove species in each listed plant genus and family.{{cite book | last=Tomlinson | first=P. B. | title=The botany of mangroves | publisher=Cambridge University Press | publication-place=Cambridge, United Kingdom | year=2016 | isbn=978-1-107-08067-6 | oclc=946579968}} Mangrove environments in the Eastern Hemisphere harbor six times as many species of trees and shrubs as do mangroves in the New World. Genetic divergence of mangrove lineages from terrestrial relatives, in combination with fossil evidence, suggests mangrove diversity is limited by evolutionary transition into the stressful marine environment, and the number of mangrove lineages has increased steadily over the Tertiary with little global extinction.{{cite journal|author=Ricklefs, R. E. |author2=A. Schwarzbach |author3=S. S. Renner |date=2006 |title=Rate of lineage origin explains the diversity anomaly in the world's mangrove vegetation |journal=American Naturalist |volume=168 |issue=6 |pages=805–810 |doi=10.1086/508711 |pmid=17109322 |s2cid=1493815 |url=http://blue.utb.edu/aschwarzbach/publications/MangroveAmerNaturalist.pdf |url-status=dead |archive-url=https://web.archive.org/web/20130616034200/http://blue.utb.edu/aschwarzbach/publications/MangroveAmerNaturalist.pdf |archive-date=16 June 2013 }}

= True mangroves =

class="wikitable mw-collapsible"
colspan=5 width=700px align=left style="background:#ddf8f8" | True mangroves (major components or strict mangroves)
colspan=5 align=center style=background:#e8e8ff | Following Tomlinson, 2016, the following 35 species are the true mangroves, contained in 5 families and 9 genera{{rp|29–30}}
Included on green backgrounds are annotations about the genera made by Tomlinson
Family

! Genus

! Mangrove species

! Common name

!

rowspan=2| Arecaceae

| colspan=4 style=background:#ddf8f8 | Monotypic subfamily within the family

Nypa

| Nypa fruticans

| Mangrove palm

| 60px

rowspan=10 style=float:top | Avicenniaceae
(disputed)

| colspan=4 style=background:#ddf8f8 | Old monogeneric family, now subsumed in Acanthaceae, but clearly isolated

rowspan=9 align=top | Avicennia

| Avicennia alba

|

| 60px

Avicennia balanophora

|

|

Avicennia bicolor

|

|

Avicennia integra

|

|

Avicennia marina

| grey mangrove
(subspecies: australasica,
eucalyptifolia, rumphiana)

| 60px

Avicennia officinalis

| Indian mangrove

| 60px

Avicennia germinans

| black mangrove

| 60px

Avicennia schaueriana

|

| 60px

Avicennia tonduzii

|

|

rowspan=4| Combretaceae

| colspan=4 style=background:#ddf8f8 | Tribe Lagunculariae (including Macropteranthes = non-mangrove)

Laguncularia

| Laguncularia racemosa

| white mangrove

| 60px

rowspan= 2 | Lumnitzera

| Lumnitzera racemosa

| white-flowered black mangrove

| 60px

Lumnitzera littorea

|

| 60px

rowspan=19 | Rhizophoraceae

| colspan=4 style=background:#ddf8f8 | Rhizophoraceae collectively form the tribe Rhizophorae, a monotypic group, within the otherwise terrestrial family

rowspan=6 | Bruguiera

| Bruguiera cylindrica

|

| 60px

Bruguiera exaristata

| rib-fruited mangrove

| 60px

Bruguiera gymnorhiza

| oriental mangrove

| 60px

Bruguiera hainesii

|

|

Bruguiera parviflora

|

| 60px

Bruguiera sexangula

| upriver orange mangrove

| 60px

rowspan=2 | Ceriops

| Ceriops australis

| yellow mangrove

| 60px

Ceriops tagal

| spurred mangrove

| 60px

rowspan=2 | Kandelia

| Kandelia candel

|

| 60px

Kandelia obovata

|

| 60px

rowspan=8 | Rhizophora

| Rhizophora apiculata

|

|

Rhizophora harrisonii

|

|

Rhizophora mangle

| red mangrove

|

Rhizophora mucronata

| Asiatic mangrove

| 60px

Rhizophora racemosa

|

|

Rhizophora samoensis

| Samoan mangrove

|

Rhizophora stylosa

| spotted mangrove,

|

Rhizophora x lamarckii

|

|

rowspan=5 | Lythraceae

| rowspan=5 | Sonneratia

| Sonneratia alba

|

| 60px

Sonneratia apetala

|

|

Sonneratia caseolaris

|

|

Sonneratia ovata

|

|

Sonneratia griffithii

|

= Other mangroves =

class="wikitable mw-collapsible"
colspan=5 width=700px align=left style="background:#ddf8f8" | Minor components
colspan=5 align=center style=background:#e8e8ff | Tomlinson, 2016, lists about 19 species as minor mangrove components, contained in 10 families and 11 genera{{rp|29–30}}
Included on green backgrounds are annotations about the genera made by Tomlinson
Family

! Genus

! Species

! Common name

!

rowspan=2| Euphorbiaceae

| colspan=4 style=background:#ddf8f8 | This genus includes about 35 non-mangrove taxa

Excoecaria

| Excoecaria agallocha

| milky mangrove, blind-your-eye mangrove and river poison tree

| 60px

rowspan=2| Lythraceae

| colspan=4 style=background:#ddf8f8 | Genus distinct in the family

Pemphis

| Pemphis acidula

| bantigue or mentigi

| 60px

rowspan=3| Malvaceae

| colspan=4 style=background:#ddf8f8 | Formerly in Bombacaceae, now an isolated genus in subfamily Bombacoideeae

rowspan=2 | Camptostemon

| Camptostemon schultzii

| kapok mangrove

| 60px

Camptostemon philippinense

|

| 60px

rowspan=3| Meliaceae

| colspan=4 style=background:#ddf8f8 | Genus of 3 species, one non-mangrove, forms tribe Xylocarpaeae with Carapa, a non–mangrove

rowspan=2 | Xylocarpus

| Xylocarpus granatum

|

| 60px

Xylocarpus moluccensis

|

| 60px

rowspan=2| Myrtaceae

| colspan=4 style=background:#ddf8f8 | An isolated genus in the family

Osbornia

| Osbornia octodonta

| mangrove myrtle

| 60px

rowspan=2| Pellicieraceae

| colspan=4 style=background:#ddf8f8 | Monotypic genus and family of uncertain phylogenetic position

Pelliciera

| Pelliciera rhizophorae

| tea mangrove

| 60px

rowspan=3| Plumbaginaceae

| colspan=4 style=background:#ddf8f8 | Isolated genus, at times segregated as family Aegialitidaceae

rowspan=2 | Aegialitis

| Aegialitis annulata

| club mangrove

| 60px

Aegialitis rotundifolia

|

| 60px

rowspan=3 | Primulaceae

| colspan=4 style=background:#ddf8f8 | Formerly an isolated genus in Myrsinaceae

rowspan=2 | Aegiceras

| Aegiceras corniculatum

| black mangrove, river mangrove or khalsi

| 60px

Aegiceras floridum

|

|

rowspan=3 | Pteridaceae

| colspan=4 style=background:#ddf8f8 | A fern somewhat isolated in its family

rowspan=2 | Acrostichum

| Acrostichum aureum

| golden leather fern, swamp fern or mangrove fern

| 60px

Acrostichum speciosum

| mangrove fern

| 60px

rowspan=2| Rubiaceae

| colspan=4 style=background:#ddf8f8 | A genus isolated in the family

Scyphiphora

| Scyphiphora hydrophylacea

| nilad

| 60px

Species distribution

{{see also|Mangrove tree distribution}}

File:Global distribution of native mangrove species.png 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].
Colour-coded number ranges indicate number of species.
Not shown are introduced ranges: Rhizophora stylosa in French Polynesia, Bruguiera sexangula, Conocarpus erectus, and Rhizophora mangle in Hawaii, Sonneratia apelata in China, and Nypa fruticans in Cameroon and Nigeria.]]

Mangroves are a type of tropical vegetation with some outliers established in subtropical latitudes, notably in South Florida and southern Japan, as well as South Africa, New Zealand and Victoria (Australia). These outliers result either from unbroken coastlines and island chains or from reliable supplies of propagules floating on warm ocean currents from rich mangrove regions.{{rp|57}}

File:Indonesia Mangrove Distribution.png

File:Global distribution of threatened mangrove species.png

{{clear left}}

"At the limits of distribution, the formation is represented by scrubby, usually monotypic Avicennia-dominated vegetation, as at Westonport Bay and Corner Inlet, Victoria, Australia. The latter locality is the highest latitude (38° 45'S) at which mangroves occur naturally. The mangroves in New Zealand, which extend as far south as 37°, are of the same type; they start as low forest in the northern part of the North Island but become low scrub toward their southern limit. In both instances, the species is referred to as Avicennia marina var. australis, although genetic comparison is clearly needed. In Western Australia, A. marina extends as far south as Bunbury (33° 19'S). In the northern hemisphere, scrubby Avicennia gerrninans in Florida occurs as far north as St. Augustine on the east coast and Cedar Point on the west. There are records of A. germinans and Rhizophora mangle for Bermuda, presumably supplied by the Gulf Stream. In southern Japan, Kandelia obovata occurs to about 31 °N (Tagawa in Hosakawa et al., 1977, but initially referred to as K. candel)."{{rp|57}}

Mangrove forests

File:Mangrove.png

{{main|Mangrove forest}}

Mangrove forests, also called mangrove swamps or mangals, are found in tropical and subtropical tidal areas. Areas where mangroves occur include estuaries and marine shorelines.

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.

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.

About 110 species are considered mangroves, in the sense of being trees that grow in such a saline swamp,{{cite web |author=Mathias, M. E. |title=Mangal (Mangrove). World Vegetation |url=http://www.botgard.ucla.edu/html/botanytextbooks/worldvegetation/marinewetlands/mangal/index.html |website=Botanical Garden, University of California at Los Angeles |publisher=Botgard.ucla.edu |access-date=8 February 2012 |url-status=dead |archive-url=https://web.archive.org/web/20120209010910/http://www.botgard.ucla.edu/html/botanytextbooks/worldvegetation/marinewetlands/mangal/index.html |archive-date=9 February 2012}} 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.{{Cite journal |last1=Sievers|first1=M. |last2=Brown|first2=C. J.|last3=Tulloch|first3=V. J. D.|last4=Pearson|first4=R. M. |last5=Haig |first5=J. A. |last6=Turschwell|first6=M. P.|last7=Connolly|first7=R. M.|date=2019|title=The Role of Vegetated Coastal Wetlands for Marine Megafauna Conservation |journal=Trends in Ecology & Evolution |volume=34 |issue=9|pages=807–817|doi=10.1016/j.tree.2019.04.004|pmid=31126633 |bibcode=2019TEcoE..34..807S |hdl=10072/391960 |s2cid=164219103 |hdl-access=free}}

File:Mangroves.jpg

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.{{cite journal |last1=Cannicci |first1=S. |last2=Fusi |first2=M. |last3=Cimó |first3=F. |last4=Dahdouh-Guebas |first4=F. |last5=Fratini |first5=S. |title=Interference competition as a key determinant for spatial distribution of mangrove crabs |journal=BMC Ecology |date=2018 |volume=18 |issue=1 |pages=8 |doi=10.1186/s12898-018-0164-1 |pmid=29448932 |pmc=5815208 |doi-access=free |bibcode=2018BMCE...18....8C }}

File:Nipa palms.jpg, the only palm species fully adapted to the mangrove biome]]

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.{{cite journal |last1=Saenger |first1=P. |last2=McConchie |first2=D. |title=Heavy metals in mangroves: methodology, monitoring and management |journal=Envis Forest Bulletin |date=2004 |volume=4 |pages=52–62 |citeseerx=10.1.1.961.9649 }}

Mangrove swamps protect coastal areas from erosion, storm surge (especially during tropical cyclones), and tsunamis.{{cite journal|last1=Mazda |first1=Y. |last2=Kobashi |first2=D. |last3=Okada |first3=S.|year=2005|title=Tidal-Scale Hydrodynamics within Mangrove Swamps |journal=Wetlands Ecology and Management |volume=13 |issue= 6 |pages=647–655 |doi=10.1007/s11273-005-0613-4 |bibcode=2005WetEM..13..647M |citeseerx=10.1.1.522.5345|s2cid=35322400}}{{cite journal |title=The Asian Tsunami: A Protective Role for Coastal Vegetation |doi=10.1126/science.1118387 |pmid=16254180 |journal=Science |volume=310 |issue=5748 |page=643 |year=2005 |last1=Danielsen |first1=F. |last2=Sørensen |first2=M. K. |last3=Olwig |first3=M. F. |last4=Selvam |first4=V. |last5=Parish |first5=F. |last6=Burgess |first6=N. D. |last7=Hiraishi |first7=T. |last8=Karunagaran |first8=V. M. |last9=Rasmussen |first9=M. S. |last10=Hansen |first10=L. B. |last11=Quarto |first11=A. |last12=Suryadiputra |first12=N. |s2cid=31945341 }}{{cite journal |title=Mangrove forest against dyke-break-induced tsunami on rapidly subsiding coasts |journal=Natural Hazards and Earth System Sciences |volume=16 |issue=7 |pages=1629–1638 |date=2016 |doi=10.5194/nhess-16-1629-2016 |last1=Takagi |first1=H. |last2=Mikami |first2=T. |last3=Fujii |first3=D. |last4=Esteban |first4=M. |last5=Kurobe |first5=S.|bibcode=2016NHESS..16.1629T |doi-access=free}} They limit high-energy wave erosion mainly during events such as storm surges and tsunamis.{{cite journal |title=How effective were mangroves as a defence against the recent tsunami? |doi=10.1016/j.cub.2005.06.008 |pmid=15964259 |journal=Current Biology |volume=15 |issue=12 |pages=R443–447 |year=2005 |last1=Dahdouh-Guebas |first1=F. |last2=Jayatissa |first2=L. P. |last3=Di Nitto |first3=D. |last4=Bosire |first4=J. O. |last5=Lo Seen |first5=D. |last6=Koedam |first6=N.|s2cid=8772526 |url=http://agritrop.cirad.fr/529549/|doi-access=free }}

The mangroves' massive root systems are efficient at dissipating wave energy.{{cite journal |doi=10.1016/s0169-5983(98)00024-0 |title=Surface wave propagation in mangrove forests |journal=Fluid Dynamics Research |volume=24 |issue=4 |pages=219 |year=1999 |last1=Massel |first1=S. R. |last2=Furukawa |first2=K. |last3=Brinkman |first3=R. M. |bibcode=1999FlDyR..24..219M|s2cid=122572658}} Likewise, they slow down tidal water so that its sediment is deposited as the tide comes in, leaving all except fine particles when the tide ebbs.{{cite journal |doi=10.1023/A:1009949411068 |year=1997 |last1=Mazda |first1=Y. |journal=Mangroves and Salt Marshes |title=Drag force due to vegetation in mangrove swamps|volume=1 |issue=3 |pages=193 |last2=Wolanski |first2=E. |last3=King |first3=B. |last4=Sase |first4=A. |last5=Ohtsuka |first5=D. |last6=Magi |first6=M.|s2cid=126945589}} In this way, mangroves build their environments. Because of the uniqueness of mangrove ecosystems and the protection against erosion they provide, they are often the object of conservation programs, including national biodiversity action plans.

The unique ecosystem found in the intricate mesh of mangrove roots offers a quiet marine habitat for young organisms.{{cite journal |doi=10.1007/s00227-010-1588-0 |pmid=24391259 |pmc=3873073 |title=Ontogenetic habitat shift, population growth, and burrowing behavior of the Indo-Pacific beach star, Archaster typicus (Echinodermata; Asteroidea) |journal=Marine Biology |volume=158 |issue=3 |pages=639–648 |year=2010 |last1=Bos |first1=A. R. |last2=Gumanao |first2=G. S. |last3=Van Katwijk |first3=M. M. |last4=Mueller |first4=B. |last5=Saceda |first5=M. M. |last6=Tejada |first6=R. L.}} 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.Encarta Encyclopedia 2005. "Seashore", by Heidi Nepf. Mangrove crabs eat the mangrove leaves, adding nutrients to the mangal mud for other bottom feeders.{{cite journal |doi=10.1007/s00442-001-0847-7 |pmid=28547499 |bibcode=2002Oecol.131....1S |title=Paradoxical selective feeding on a low-nutrient diet: Why do mangrove crabs eat leaves? |journal=Oecologia |volume=131 |issue=1 |pages=1–7 |year=2002 |last1=Skov |first1=M. W. |last2=Hartnoll |first2=R.G.|s2cid=23407273}} In at least some cases, the export of carbon fixed in mangroves is important in coastal food webs.{{Cite journal|author1=Abrantes, K. G. |author2=Johnston, R. |author3=Connolly, R. M. |author4=Sheaves, M. |date=2015 |title=Importance of Mangrove Carbon for Aquatic Food Webs in Wet–Dry Tropical Estuaries |journal=Estuaries and Coasts |volume=38 |issue=1|pages=383–399 |doi=10.1007/s12237-014-9817-2 |bibcode=2015EstCo..38..383A |hdl=10072/141734|s2cid=3957868|issn=1559-2731|hdl-access=free}}

Mangrove forests contribute significantly to coastal ecosystems by fostering complex and diverse food webs. The intricate root systems of mangroves create a habitat conducive to the proliferation of microorganisms, crustaceans, and small fish, forming the foundational tiers of the food chain. This abundance of organisms serves as a critical food source for larger predators like birds, reptiles, and mammals within the ecosystem. Additionally, mangrove forests function as essential nurseries for many commercially important fish species, providing a sheltered environment rich in nutrients during their early life stages. The decomposition of leaves and organic matter in the water further enhances the nutrient content, supporting overall ecosystem productivity. In summary, mangrove forests play a crucial and unbiased role in sustaining biodiversity and ecological balance within coastal food webs.{{Cite journal |last1=Muro-Torres |first1=Victor M. |last2=Amezcua |first2=Felipe |last3=Soto-Jiménez |first3=Martin |last4=Balart |first4=Eduardo F. |last5=Serviere-Zaragoza |first5=Elisa |last6=Green |first6=Lucinda |last7=Rajnohova |first7=Jana |date=2020-11-05 |title=Primary Sources and Food Web Structure of a Tropical Wetland with High Density of Mangrove Forest |journal=Water |language=en |volume=12 |issue=11 |pages=3105 |doi=10.3390/w12113105 |doi-access=free |issn=2073-4441|hdl=1854/LU-01HV3XGJPZJE3Z72394VV0MRJB |hdl-access=free }}

Larger marine organisms benefit from the habitat as a nursery for their offspring. Lemon sharks depend on mangrove creeks to give birth to their pups. The ecosystem provides little competition and minimizes threats of predation to juvenile lemon sharks as they use the cover of mangroves to practice hunting before entering the food web of the ocean.{{Cite journal |last1=Newman |first1=Sp |last2=Handy |first2=Rd |last3=Gruber |first3=Sh |date=2010-01-05 |title=Diet and prey preference of juvenile lemon sharks Negaprion brevirostris |url=http://www.int-res.com/abstracts/meps/v398/p221-234/ |journal=Marine Ecology Progress Series |language=en |volume=398 |pages=221–234 |doi=10.3354/meps08334 |bibcode=2010MEPS..398..221N |issn=0171-8630}}

Mangrove plantations in Vietnam, Thailand, Philippines, and India host several commercially important species of fish and crustaceans.{{Cite book|last1=Gupta|first1=S. K.|url=https://books.google.com/books?id=KuneDwAAQBAJ&pg=PA34 |title=Soil Salinity Management in Agriculture: Technological Advances and Applications |last2=Goyal |first2=M. R. |date=2017|publisher=CRC Press|isbn=978-1-315-34177-4}}

The mangrove food chain extends beyond the marine ecosystem. Coastal bird species inhabit the tidal ecosystems feeding off small marine organisms and wetland insects. Common bird families found in mangroves around the world are egrets, kingfishers, herons, and hornbills, among many others dependent on ecological range.{{Cite journal |last1=Mohd-Taib |first1=Farah Shafawati |last2=Mohd-Saleh |first2=Wardah |last3=Asyikha |first3=Rosha |last4=Mansor |first4=Mohammad Saiful |last5=Ahmad-Mustapha |first5=Muzzneena |last6=Mustafa-Bakray |first6=Nur Aqilah |last7=Mod-Husin |first7=Shahril |last8=Md-Shukor |first8=Aisah |last9=Amat-Darbis |first9=Nurul Darsani |last10=Sulaiman |first10=Norela |date=June 2020 |title=Effects of anthropogenic disturbance on the species assemblages of birds in the back mangrove forests |url=https://link.springer.com/10.1007/s11273-020-09726-z |journal=Wetlands Ecology and Management |language=en |volume=28 |issue=3 |pages=479–494 |doi=10.1007/s11273-020-09726-z |bibcode=2020WetEM..28..479M |s2cid=218484236 |issn=0923-4861}} Bird predation plays a key role in maintaining prey species along coastlines and within mangrove ecosystems.

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.{{cite journal |doi=10.1002/ggge.20194 |title=Degradation of mangrove tissues by arboreal termites (Nasutitermes acajutlae) and their role in the mangrove C cycle (Puerto Rico): Chemical characterization and organic matter provenance using bulk δ13C, C/N, alkaline CuO oxidation-GC/MS, and solid-state |journal=Geochemistry, Geophysics, Geosystems |volume=14 |issue=8 |page=3176 |year=2013 |last1=Vane |first1=C. H. |last2=Kim |first2=A. W. |last3=Moss-Hayes |first3=V. |last4=Snape |first4=C. E. |last5=Diaz |first5=M. C. |last6=Khan |first6=N. S. |last7=Engelhart |first7=S. E. |last8=Horton |first8=B. P. |bibcode=2013GGG....14.3176V |doi-access=free}} 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).

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. Mangrove forests are an important part of the cycling and storage of carbon in tropical coastal ecosystems. Knowing this, scientists seek to reconstruct the environment and investigate changes to the coastal ecosystem over thousands of years using sediment cores.{{cite journal |last1=Versteegh |first1=G.J. |display-authors=et al. |year=2004 |title=Taraxerol and Rhizophora pollen as proxies for tracking past mangrove ecosystems |journal=Geochimica et Cosmochimica Acta |volume=68 |issue=3 |pages=411–22 |doi=10.1016/S0016-7037(03)00456-3 |bibcode=2004GeCoA..68..411V}} 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. Termites play an important role in the formation of peat from mangrove materials. They process fallen leaf litter, root systems and wood from mangroves into peat to build their nests, and stabilise the chemistry of this peat that represents approximately 2% of above ground carbon storage in mangroves. As the nests are buried over time this carbon is stored in the sediment and the carbon cycle continues.

Mangroves are an important source of blue carbon. Globally, mangroves stored {{Cvt|4.19|Gt|lb|abbr=unit}} of carbon in 2012. Two percent of global mangrove carbon was lost between 2000 and 2012, equivalent to a maximum potential of {{Cvt|0.316996250|Gt|lbs}} of emissions of carbon dioxide in Earth's atmosphere.{{Cite journal|last1=Hamilton |first1=S. E. |last2=Friess |first2=D. A. |date=2018 |title=Global carbon stocks and potential emissions due to mangrove deforestation from 2000 to 2012 |journal=Nature Climate Change |volume=8 |issue=3|pages=240–244 |doi=10.1038/s41558-018-0090-4 |bibcode=2018NatCC...8..240H|arxiv=1611.00307 |s2cid=89785740}}

Globally, mangroves have been shown to provide measurable economic protections to coastal communities affected by tropical storms.{{Cite journal|last1=Hochard|first1=J. P. |last2=Hamilton |first2=S. |last3=Barbier |first3=E. B.|date=2019 |title=Mangroves shelter coastal economic activity from cyclones |journal=Proceedings of the National Academy of Sciences|volume=116|issue=25|pages=12232–12237 |doi-access=free |doi=10.1073/pnas.1820067116|pmid=31160457 |pmc=6589649|bibcode=2019PNAS..11612232H}}

Mangrove microbiome

{{see also|Plant microbiome}}

Plant microbiomes play crucial roles in the health and productivity of mangroves.{{cite journal |doi = 10.3389/fpls.2018.01563|doi-access = free|title = Plant Microbiome and Its Link to Plant Health: Host Species, Organs and Pseudomonas syringae pv. Actinidiae Infection Shaping Bacterial Phyllosphere Communities of Kiwifruit Plants|year = 2018|last1 = Purahong|first1 = Witoon|last2 = Orrù|first2 = Luigi|last3 = Donati|first3 = Irene|last4 = Perpetuini|first4 = Giorgia|last5 = Cellini|first5 = Antonio|last6 = Lamontanara|first6 = Antonella|last7 = Michelotti|first7 = Vania|last8 = Tacconi|first8 = Gianni|last9 = Spinelli|first9 = Francesco|journal = Frontiers in Plant Science|volume = 9|page = 1563|pmid = 30464766|pmc = 6234494}} Many researchers have successfully applied knowledge acquired about plant microbiomes to produce specific inocula for crop protection.{{Cite journal |last1=Afzal |first1=A. |last2=Bano |first2=A. |date=2008 |title=Rhizobium and phosphate solubilizing bacteria improve the yield and phosphorus uptake in wheat (Triticum aestivum) |url=https://www.researchgate.net/publication/228684375 |journal=International Journal of Agriculture and Biology (Pakistan) |volume=10 |issue=1 |pages=85–88 |issn=1560-8530 |eissn=1814-9596}} Such inocula can stimulate plant growth by releasing phytohormones and enhancing uptake of some mineral nutrients (particularly phosphorus and nitrogen).{{cite journal |doi = 10.1371/journal.pbio.2001793|title = Research priorities for harnessing plant microbiomes in sustainable agriculture|year = 2017|last1 = Busby|first1 = Posy E.|last2 = Soman|first2 = Chinmay|last3 = Wagner|first3 = Maggie R.|last4 = Friesen|first4 = Maren L.|last5 = Kremer|first5 = James|last6 = Bennett|first6 = Alison|last7 = Morsy|first7 = Mustafa|last8 = Eisen|first8 = Jonathan A.|last9 = Leach|first9 = Jan E.|last10 = Dangl|first10 = Jeffery L.|journal = PLOS Biology|volume = 15|issue = 3|pages = e2001793|pmid = 28350798|pmc = 5370116 | doi-access=free }}{{cite journal |doi = 10.1016/j.tplants.2012.04.001|title = The rhizosphere microbiome and plant health|year = 2012|last1 = Berendsen|first1 = Roeland L.|last2 = Pieterse|first2 = Corné M.J.|last3 = Bakker|first3 = Peter A. H. M.|journal = Trends in Plant Science|volume = 17|issue = 8|pages = 478–486|pmid = 22564542| bibcode=2012TPS....17..478B |hdl = 1874/255269| s2cid=32900768 |hdl-access = free}}{{Cite journal |last1=Bringel |first1=Françoise |last2=Couée |first2=Ivan |year=2015 |title=Pivotal roles of phyllosphere microorganisms at the interface between plant functioning and atmospheric trace gas dynamics |journal=Frontiers in Microbiology |volume=06 |page=486 |doi=10.3389/fmicb.2015.00486 |pmc=4440916 |pmid=26052316 |doi-access=free}} However, most of the plant microbiome studies have focused on the model plant Arabidopsis thaliana and economically important crop plants, such as rice, barley, wheat, maize and soybean. There is less information on the microbiomes of tree species. Plant microbiomes are determined by plant-related factors (e.g., genotype, organ, species, and health status) and environmental factors (e.g., land use, climate, and nutrient availability). Two of the plant-related factors, plant species and genotypes, have been shown to play significant roles in shaping rhizosphere and plant microbiomes, as tree genotypes and species are associated with specific microbial communities. Different plant organs also have specific microbial communities depending on plant-associated factors (plant genotype, available nutrients, and organ-specific physicochemical conditions) and environmental conditions (associated with aboveground and underground surfaces and disturbances).{{cite journal |doi = 10.1111/nph.13697|title = Plant compartment and biogeography affect microbiome composition in cultivated and native Agave species|year = 2016|last1 = Coleman-Derr|first1 = Devin|last2 = Desgarennes|first2 = Damaris|last3 = Fonseca-Garcia|first3 = Citlali|last4 = Gross|first4 = Stephen|last5 = Clingenpeel|first5 = Scott|last6 = Woyke|first6 = Tanja|last7 = North|first7 = Gretchen|last8 = Visel|first8 = Axel|last9 = Partida-Martinez|first9 = Laila P.|last10 = Tringe|first10 = Susannah G.|journal = New Phytologist|volume = 209|issue = 2|pages = 798–811|pmid = 26467257|pmc = 5057366}}{{cite journal |doi = 10.1186/s40168-018-0413-8|title = The Populus holobiont: Dissecting the effects of plant niches and genotype on the microbiome|year = 2018|last1 = Cregger|first1 = M. A.|last2 = Veach|first2 = A. M.|last3 = Yang|first3 = Z. K.|last4 = Crouch|first4 = M. J.|last5 = Vilgalys|first5 = R.|last6 = Tuskan|first6 = G. A.|last7 = Schadt|first7 = C. W.|journal = Microbiome|volume = 6|issue = 1|page = 31|pmid = 29433554|pmc = 5810025 | doi-access=free }}{{cite journal |doi = 10.1111/nph.13760|title = Disentangling the factors shaping microbiota composition across the plant holobiont|year = 2016|last1 = Hacquard|first1 = Stéphane|journal = New Phytologist|volume = 209|issue = 2|pages = 454–457|pmid = 26763678| hdl=11858/00-001M-0000-002B-166F-5 |doi-access = free}}

=Root microbiome=

File:Bacterial and fungal community in a mangrove tree.webp 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]. Bacterial taxonomic community composition in the rhizosphere soil and fungal taxonomic community composition in all four rhizosphere soil and plant compartments. Information on the fungal ecological functional groups is also provided. Proportions of fungal OTUs (approximate species) that can colonise at least two of the compartments are shown in the left panel.]]

{{see also|Root microbiome}}

Mangrove roots harbour a repertoire of microbial taxa that contribute to important ecological functions in mangrove ecosystems. Like typical terrestrial plants, mangroves depend upon mutually beneficial interactions with microbial communities.{{cite journal |doi = 10.1007/s13213-012-0442-7|title = Biodiversity and biotechnological potential of microorganisms from mangrove ecosystems: A review|year = 2013|last1 = Thatoi|first1 = Hrudayanath|last2 = Behera|first2 = Bikash Chandra|last3 = Mishra|first3 = Rashmi Ranjan|last4 = Dutta|first4 = Sushil Kumar|journal = Annals of Microbiology|volume = 63|pages = 1–19|s2cid = 17798850|doi-access = free}} In particular, microbes residing in developed roots could help mangroves transform nutrients into usable forms before plant assimilation.{{cite journal |doi = 10.1016/j.scitotenv.2020.137807|title = Revealing structure and assembly for rhizophyte-endophyte diazotrophic community in mangrove ecosystem after introduced Sonneratia apetala and Laguncularia racemosa|year = 2020|last1 = Liu|first1 = Xingyu|last2 = Yang|first2 = Chao|last3 = Yu|first3 = Xiaoli|last4 = Yu|first4 = Huang|last5 = Zhuang|first5 = Wei|last6 = Gu|first6 = Hang|last7 = Xu|first7 = Kui|last8 = Zheng|first8 = Xiafei|last9 = Wang|first9 = Cheng|last10 = Xiao|first10 = Fanshu|last11 = Wu|first11 = Bo|last12 = He|first12 = Zhili|last13 = Yan|first13 = Qingyun|journal = Science of the Total Environment|volume = 721|page = 137807|pmid = 32179356|bibcode = 2020ScTEn.72137807L|s2cid = 212739128}}{{cite journal |doi = 10.1038/s41467-018-07343-2|title = The structure and function of the global citrus rhizosphere microbiome|year = 2018|last1 = Xu|first1 = Jin|last2 = Zhang|first2 = Yunzeng|last3 = Zhang|first3 = Pengfan|last4 = Trivedi|first4 = Pankaj|last5 = Riera|first5 = Nadia|last6 = Wang|first6 = Yayu|last7 = Liu|first7 = Xin|last8 = Fan|first8 = Guangyi|last9 = Tang|first9 = Jiliang|last10 = Coletta-Filho|first10 = Helvécio D.|last11 = Cubero|first11 = Jaime|last12 = Deng|first12 = Xiaoling|last13 = Ancona|first13 = Veronica|last14 = Lu|first14 = Zhanjun|last15 = Zhong|first15 = Balian|last16 = Roper|first16 = M. Caroline|last17 = Capote|first17 = Nieves|last18 = Catara|first18 = Vittoria|last19 = Pietersen|first19 = Gerhard|last20 = Vernière|first20 = Christian|last21 = Al-Sadi|first21 = Abdullah M.|last22 = Li|first22 = Lei|last23 = Yang|first23 = Fan|last24 = Xu|first24 = Xun|last25 = Wang|first25 = Jian|last26 = Yang|first26 = Huanming|last27 = Jin|first27 = Tao|last28 = Wang|first28 = Nian|journal = Nature Communications|volume = 9|issue = 1|page = 4894|pmid = 30459421|pmc = 6244077|bibcode = 2018NatCo...9.4894X}} These microbes also provide mangroves phytohormones for suppressing phytopathogens or helping mangroves withstand heat and salinity. In turn, root-associated microbes receive carbon metabolites from the plant via root exudates,{{cite journal | last1=Sasse | first1=Joelle | last2=Martinoia | first2=Enrico | last3=Northen | first3=Trent | title=Feed Your Friends: Do Plant Exudates Shape the Root Microbiome? | journal=Trends in Plant Science | publisher=Elsevier BV | volume=23 | issue=1 | year=2018 | issn=1360-1385 | doi=10.1016/j.tplants.2017.09.003 | pages=25–41| pmid=29050989 | bibcode=2018TPS....23...25S | osti=1532289 | s2cid=205455681 | url=https://www.zora.uzh.ch/id/eprint/148899/1/Sasse_TIPS_2017.pdf }} thus close associations between the plant and microbes are established for their mutual benefits.{{cite journal |doi = 10.1146/annurev.arplant.57.032905.105159|title = The Role of Root Exudates in Rhizosphere Interactions with Plants and Other Organisms|year = 2006|last1 = Bais|first1 = Harsh P.|last2 = Weir|first2 = Tiffany L.|last3 = Perry|first3 = Laura G.|last4 = Gilroy|first4 = Simon|last5 = Vivanco|first5 = Jorge M.|journal = Annual Review of Plant Biology|volume = 57|pages = 233–266|pmid = 16669762}}{{cite journal |doi = 10.1038/s41522-020-00164-6|title = Diversity, function and assembly of mangrove root-associated microbial communities at a continuous fine-scale|year = 2020|last1 = Zhuang|first1 = Wei|last2 = Yu|first2 = Xiaoli|last3 = Hu|first3 = Ruiwen|last4 = Luo|first4 = Zhiwen|last5 = Liu|first5 = Xingyu|last6 = Zheng|first6 = Xiafei|last7 = Xiao|first7 = Fanshu|last8 = Peng|first8 = Yisheng|last9 = He|first9 = Qiang|last10 = Tian|first10 = Yun|last11 = Yang|first11 = Tony|last12 = Wang|first12 = Shanquan|last13 = Shu|first13 = Longfei|last14 = Yan|first14 = Qingyun|last15 = Wang|first15 = Cheng|last16 = He|first16 = Zhili|journal = npj Biofilms and Microbiomes|volume = 6|issue = 1|page = 52|pmid = 33184266|pmc = 7665043}} 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].

The taxonomic class level shows that most Proteobacteria were reported to come from Gammaproteobacteria, followed by Deltaproteobacteria and Alphaproteobacteria. The diverse function and the phylogenic variation of Gammaproteobacteria, which consisted of orders such as Alteromonadales and Vibrionales, are found in marine and coastal regions and are high in abundance in mangrove sediments functioning as nutrient recyclers. Members of Deltaproteobacteria found in mangrove soil are mostly sulfur-related, consisting of Desulfobacterales, Desulfuromonadales, Desulfovibrionales, and Desulfarculales among others.{{cite journal |last1=Lai |first1=Jiayong |last2=Cheah |first2=Wee |last3=Palaniveloo |first3=Kishneth |last4=Suwa |first4=Rempei |last5=Sharma |first5=Sahadev |title=A Systematic Review of the Physicochemical and Microbial Diversity of Well-Preserved, Restored, and Disturbed Mangrove Forests: What Is Known and What Is the Way Forward? |journal=Forests |date=16 December 2022 |volume=13 |issue=12 |pages=2160 |doi=10.3390/f13122160 |doi-access=free }}

Highly diverse microbial communities (mainly bacteria and fungi) have been found to inhabit and function in mangrove roots.{{cite journal |doi = 10.1007/s00468-015-1233-0|title = Mangrove root: Adaptations and ecological importance|year = 2016|last1 = Srikanth|first1 = Sandhya|last2 = Lum|first2 = Shawn Kaihekulani Yamauchi|last3 = Chen|first3 = Zhong|journal = Trees|volume = 30|issue = 2|pages = 451–465| bibcode=2016Trees..30..451S |s2cid = 5471541}}{{cite journal |doi = 10.2307/2261526|jstor = 2261526|title = Soil Physicochemical Patterns and Mangrove Species Distribution--Reciprocal Effects?|last1 = McKee|first1 = Karen L.|journal = Journal of Ecology|year = 1993|volume = 81|issue = 3|pages = 477–487| bibcode=1993JEcol..81..477M }} For example, diazotrophic bacteria in the vicinity of mangrove roots could perform biological nitrogen fixation, which provides 40–60% of the total nitrogen required by mangroves;{{cite journal |doi = 10.1007/s003740000319|title = The role of sediment microorganisms in the productivity, conservation, and rehabilitation of mangrove ecosystems: An overview|year = 2001|last1 = Holguin|first1 = Gina|last2 = Vazquez|first2 = Patricia|last3 = Bashan|first3 = Yoav|journal = Biology and Fertility of Soils|volume = 33|issue = 4|pages = 265–278| bibcode=2001BioFS..33..265H |s2cid = 10826862}}{{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|doi-access = free}} the soil attached to mangrove roots lacks oxygen but is rich in organic matter, providing an optimal microenvironment for sulfate-reducing bacteria and methanogens, ligninolytic, cellulolytic, and amylolytic fungi are prevalent in the mangrove root environment; rhizosphere fungi could help mangroves survive in waterlogged and nutrient-restricted environments.{{cite journal |doi = 10.1016/j.apsoil.2013.11.009|title = Effects of arbuscular mycorrhizal inoculation and phosphorus supply on the growth and nutrient uptake of Kandelia obovata (Sheue, Liu & Yong) seedlings in autoclaved soil|year = 2014|last1 = Xie|first1 = Xiangyu|last2 = Weng|first2 = Bosen|last3 = Cai|first3 = Bangping|last4 = Dong|first4 = Yiran|last5 = Yan|first5 = Chongling|journal = Applied Soil Ecology|volume = 75|pages = 162–171| bibcode=2014AppSE..75..162X }} These studies have provided increasing evidence to support the importance of root-associated bacteria and fungi for mangrove growth and health.

Recent studies have investigated the detailed structure of root-associated microbial communities at a continuous fine-scale in other plants,{{cite journal | last1=Edwards | first1=Joseph | last2=Johnson | first2=Cameron | last3=Santos-Medellín | first3=Christian | last4=Lurie | first4=Eugene | last5=Podishetty | first5=Natraj Kumar | last6=Bhatnagar | first6=Srijak | last7=Eisen | first7=Jonathan A. | last8=Sundaresan | first8=Venkatesan | title=Structure, variation, and assembly of the root-associated microbiomes of rice | journal=Proceedings of the National Academy of Sciences | volume=112 | issue=8 | date=20 January 2015 | issn=0027-8424 | doi=10.1073/pnas.1414592112 | pages=E911–E920| pmid=25605935 | pmc=4345613 | bibcode=2015PNAS..112E.911E | doi-access=free }} where a microhabitat was divided into four root compartments: endosphere,{{cite journal |doi = 10.1016/j.cell.2018.10.020|title = Microbial Interkingdom Interactions in Roots Promote Arabidopsis Survival|year = 2018|last1 = Durán|first1 = Paloma|last2 = Thiergart|first2 = Thorsten|last3 = Garrido-Oter|first3 = Ruben|last4 = Agler|first4 = Matthew|last5 = Kemen|first5 = Eric|last6 = Schulze-Lefert|first6 = Paul|last7 = Hacquard|first7 = Stéphane|journal = Cell|volume = 175|issue = 4|pages = 973–983.e14|pmid = 30388454|pmc = 6218654}}{{cite journal |doi = 10.1042/BCJ20180615|title = Interactions between plants and soil shaping the root microbiome under abiotic stress|year = 2019|last1 = Hartman|first1 = Kyle|last2 = Tringe|first2 = Susannah G.|journal = Biochemical Journal|volume = 476|issue = 19|pages = 2705–2724|pmid = 31654057|pmc = 6792034}} episphere, rhizosphere,{{cite journal |doi = 10.1073/pnas.1414592112|title = Structure, variation, and assembly of the root-associated microbiomes of rice|year = 2015|last1 = Edwards|first1 = Joseph|last2 = Johnson|first2 = Cameron|last3 = Santos-Medellín|first3 = Christian|last4 = Lurie|first4 = Eugene|last5 = Podishetty|first5 = Natraj Kumar|last6 = Bhatnagar|first6 = Srijak|last7 = Eisen|first7 = Jonathan A.|last8 = Sundaresan|first8 = Venkatesan|journal = Proceedings of the National Academy of Sciences|volume = 112|issue = 8|pages = E911–E920|pmid = 25605935|pmc = 4345613|bibcode = 2015PNAS..112E.911E|doi-access = free}}{{cite journal |doi = 10.1146/annurev-phyto-082712-102342|title = Roots Shaping Their Microbiome: Global Hotspots for Microbial Activity|year = 2015|last1 = Reinhold-Hurek|first1 = Barbara|last2 = Bünger|first2 = Wiebke|last3 = Burbano|first3 = Claudia Sofía|last4 = Sabale|first4 = Mugdha|last5 = Hurek|first5 = Thomas|journal = Annual Review of Phytopathology|volume = 53|pages = 403–424|pmid = 26243728}} and nonrhizosphere or bulk soil.{{cite journal |last1=Liu |first1=Yalong |last2=Ge |first2=Tida |last3=Ye |first3=Jun |last4=Liu |first4=Shoulong |last5=Shibistova |first5=Olga |last6=Wang |first6=Ping |last7=Wang |first7=Jingkuan |last8=Li |first8=Yong |last9=Guggenberger |first9=Georg |last10=Kuzyakov |first10=Yakov |author-link10=Yakov Kuzyakov |last11=Wu |first11=Jinshui |year=2019 |title=Initial utilization of rhizodeposits with rice growth in paddy soils: Rhizosphere and N fertilization effects |journal=Geoderma |volume=338 |pages=30–39 |bibcode=2019Geode.338...30L |doi=10.1016/j.geoderma.2018.11.040 |s2cid=134648694}}{{cite journal |doi = 10.1016/j.femsec.2003.11.012|title = Microbial interactions in the mycorrhizosphere and their significance for sustainable agriculture|year = 2004|last1 = Johansson|first1 = Jonas F.|last2 = Paul|first2 = Leslie R.|last3 = Finlay|first3 = Roger D.|journal = FEMS Microbiology Ecology|volume = 48|issue = 1|pages = 1–13|pmid = 19712426| s2cid=22700384 |doi-access = free| bibcode=2004FEMME..48....1J }} Moreover, the microbial communities in each compartment have been reported to have unique characteristics. Root exudates selectively enrich adapted microbial populations; however, these exudates were found to exert only marginal impacts on microbes in the bulk soil outside the rhizosphere .{{cite journal |doi = 10.1016/j.tplants.2017.09.003|title = Feed Your Friends: Do Plant Exudates Shape the Root Microbiome?|year = 2018|last1 = Sasse|first1 = Joelle|last2 = Martinoia|first2 = Enrico|last3 = Northen|first3 = Trent|journal = Trends in Plant Science|volume = 23|issue = 1|pages = 25–41|pmid = 29050989| bibcode=2018TPS....23...25S |osti = 1532289| s2cid=205455681 |url = https://www.zora.uzh.ch/id/eprint/148899/1/Sasse_TIPS_2017.pdf}} Furthermore, it was noted that the root episphere, rather than the rhizosphere, was primarily responsible for controlling the entry of specific microbial populations into the root, resulting in the selective enrichment of Proteobacteria in the endosphere.{{cite journal |doi = 10.1038/ncomms5950|title = Niche and host-associated functional signatures of the root surface microbiome|year = 2014|last1 = Ofek-Lalzar|first1 = Maya|last2 = Sela|first2 = Noa|last3 = Goldman-Voronov|first3 = Milana|last4 = Green|first4 = Stefan J.|last5 = Hadar|first5 = Yitzhak|last6 = Minz|first6 = Dror|journal = Nature Communications|volume = 5|page = 4950|pmid = 25232638|bibcode = 2014NatCo...5.4950O|doi-access = free}} These findings provide new insights into the niche differentiation of root-associated microbial communities, Nevertheless, amplicon-based community profiling may not provide the functional characteristics of root-associated microbial communities in plant growth and biogeochemical cycling.{{cite journal |doi = 10.1007/s13238-020-00724-8|title = A practical guide to amplicon and metagenomic analysis of microbiome data|year = 2021|last1 = Liu|first1 = Yong-Xin|last2 = Qin|first2 = Yuan|last3 = Chen|first3 = Tong|last4 = Lu|first4 = Meiping|last5 = Qian|first5 = Xubo|last6 = Guo|first6 = Xiaoxuan|last7 = Bai|first7 = Yang|journal = Protein & Cell|volume = 12|issue = 5|pages = 315–330|pmid = 32394199|pmc = 8106563}} Unraveling functional patterns across the four root compartments holds a great potential for understanding functional mechanisms responsible for mediating root–microbe interactions in support of enhancing mangrove ecosystem functioning.

The diversity of bacteria in disturbed mangroves is reported to be higher than in

well-preserved mangroves Studies comparing mangroves in different conservation states show that bacterial composition in disturbed mangrove sediment alters its structure, leading to a functional equilibrium, where the dynamics of chemicals in mangrove soils lead to the remodeling of its microbial structure.{{cite journal |title=Exploring bacterial functionality in mangrove sediments and its capability to overcome anthropogenic activity |date=2019 |doi=10.1016/j.marpolbul.2019.03.001 |last1=Cotta |first1=Simone Raposo |last2=Cadete |first2=Luana Lira |last3=Van Elsas |first3=Jan Dirk |last4=Andreote |first4=Fernando Dini |last5=Dias |first5=Armando Cavalcante Franco |journal=Marine Pollution Bulletin |volume=141 |pages=586–594 |pmid=30955771 |bibcode=2019MarPB.141..586C |s2cid=91872087 |url=https://research.rug.nl/en/publications/45e6976e-8216-46e2-a2bb-d23a6158b694 }}

=Suggestions for future mangrove microbial diversity research=

Despite many research advancements in mangrove sediment bacterial metagenomics

diversity in various conditions over the past few years, bridging the research gap and

expanding our knowledge towards the relationship between microbes mainly constituted of bacteria and its nutrient cycles in the mangrove sediment and direct and indirect impacts on mangrove growth and stand-structures as coastal barriers and other ecological service providers. Thus, based on studies by Lai et al.'s systematic review, here they suggest sampling improvements and a fundamental environmental index for future reference.

=Mangrove virome=

File:Caudovirales.svgs are viruses that infect bacteria, such as cyanobacteria. Shown are the virions of different families of tailed phages: Myoviridae, Podoviridae and Siphoviridae]]

{{see also|Virome|Marine viruses}}

File:Phylogenetic tree of Caudovirales from mangrove virome contigs.webp 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]. Reference sequences are coloured black, and virome contigs are indicated with varied colours. The scale bar represents half amino acid substitution per site.]]

Mangrove forests are one of the most carbon-rich biomes, accounting for 11% of the total input of terrestrial carbon into oceans. Viruses are thought to significantly influence local and global biogeochemical cycles, though as of 2019, little information was available about the community structure, genetic diversity, and ecological roles of viruses in mangrove ecosystems.

Viruses are the most abundant biological entities on earth, present in virtually all ecosystems.{{cite journal |doi = 10.1038/nature04160|title = Viruses in the sea|year = 2005|last1 = Suttle|first1 = Curtis A.|journal = Nature|volume = 437|issue = 7057|pages = 356–361|pmid = 16163346|bibcode = 2005Natur.437..356S|s2cid = 4370363}}{{cite journal |doi = 10.1073/pnas.1305956110|title = Twelve previously unknown phage genera are ubiquitous in global oceans|year = 2013|last1 = Holmfeldt|first1 = K.|last2 = Solonenko|first2 = N.|last3 = Shah|first3 = M.|last4 = Corrier|first4 = K.|last5 = Riemann|first5 = L.|last6 = Verberkmoes|first6 = N. C.|last7 = Sullivan|first7 = M. B.|journal = Proceedings of the National Academy of Sciences|volume = 110|issue = 31|pages = 12798–12803|pmid = 23858439|pmc = 3732932|bibcode = 2013PNAS..11012798H|doi-access = free}} By lysing their hosts, that is, by rupturing their cell membranes, viruses control host abundance and affect the structure of host communities.{{cite journal |doi = 10.3389/fmicb.2014.00355|doi-access = free|title = Environmental bacteriophages: Viruses of microbes in aquatic ecosystems|year = 2014|last1 = Sime-Ngando|first1 = TéLesphore|journal = Frontiers in Microbiology|volume = 5|page = 355|pmid = 25104950|pmc = 4109441}} Viruses also influence their host diversity and evolution through horizontal gene transfer, selection for resistance and manipulation of bacterial metabolisms.{{cite journal |doi = 10.1146/annurev-marine-120709-142805|title = Marine Viruses: Truth or Dare|year = 2012|last1 = Breitbart|first1 = Mya|journal = Annual Review of Marine Science|volume = 4|pages = 425–448|pmid = 22457982|bibcode = 2012ARMS....4..425B}}{{cite journal |doi = 10.1128/mBio.00893-17|title = Deep-Sea Hydrothermal Vent Viruses Compensate for Microbial Metabolism in Virus-Host Interactions|year = 2017|last1 = He|first1 = Tianliang|last2 = Li|first2 = Hongyun|last3 = Zhang|first3 = Xiaobo|journal = mBio|volume = 8|issue = 4|pmid = 28698277|pmc = 5513705}}{{cite journal |doi = 10.1073/pnas.1319778111|title = Modeling ecological drivers in marine viral communities using comparative metagenomics and network analyses|year = 2014|last1 = Hurwitz|first1 = B. L.|last2 = Westveld|first2 = A. H.|last3 = Brum|first3 = J. R.|last4 = Sullivan|first4 = M. B.|journal = Proceedings of the National Academy of Sciences|volume = 111|issue = 29|pages = 10714–10719|pmid = 25002514|pmc = 4115555|bibcode = 2014PNAS..11110714H|doi-access = free}} Importantly, marine viruses affect local and global biogeochemical cycles through the release of substantial amounts of organic carbon and nutrients from hosts and assist microbes in driving biogeochemical cycles with auxiliary metabolic genes (AMGs).{{cite journal |doi = 10.1126/science.1252229

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|s2cid = 692770

|hdl-access = free

}}{{cite journal |doi = 10.1038/nrmicro.2017.113|title = Algal virus boosts nitrogen uptake in the ocean|year = 2017|last1 = York|first1 = Ashley|journal = Nature Reviews Microbiology|volume = 15|issue = 10|page = 573|pmid = 28900307|s2cid = 19473466|doi-access = free}}{{cite journal |doi = 10.1038/nature19366|title = Ecogenomics and potential biogeochemical impacts of globally abundant ocean viruses|year = 2016|last1 = Roux|first1 = Simon|last2 = Brum|first2 = Jennifer R.|last3 = Dutilh|first3 = Bas E.|last4 = Sunagawa|first4 = Shinichi|last5 = Duhaime|first5 = Melissa B.|last6 = Loy|first6 = Alexander|last7 = Poulos|first7 = Bonnie T.|last8 = Solonenko|first8 = Natalie|last9 = Lara|first9 = Elena|last10 = Poulain|first10 = Julie|last11 = Pesant|first11 = Stéphane|last12 = Kandels-Lewis|first12 = Stefanie|last13 = Dimier|first13 = Céline|last14 = Picheral|first14 = Marc|last15 = Searson|first15 = Sarah|last16 = Cruaud|first16 = Corinne|last17 = Alberti|first17 = Adriana|last18 = Duarte|first18 = Carlos M.|last19 = Gasol|first19 = Josep M.|last20 = Vaqué|first20 = Dolors|last21 = Bork|first21 = Peer|last22 = Acinas|first22 = Silvia G.|last23 = Wincker|first23 = Patrick|last24 = Sullivan|first24 = Matthew B.|journal = Nature|volume = 537|issue = 7622|pages = 689–693|pmid = 27654921|bibcode = 2016Natur.537..689.|hdl = 1874/341494|s2cid = 54182070|hdl-access = free}}

It is presumed that AMGs augment viral-infected host metabolism and facilitate the production of new viruses.{{cite journal |doi = 10.1038/nature08060|title = Viruses manipulate the marine environment|year = 2009|last1 = Rohwer|first1 = Forest|last2 = Thurber|first2 = Rebecca Vega|journal = Nature|volume = 459|issue = 7244|pages = 207–212|pmid = 19444207|bibcode = 2009Natur.459..207R|s2cid = 4397295}} AMGs have been extensively explored in marine cyanophages and include genes involved in photosynthesis, carbon turnover, phosphate uptake and stress response.{{cite journal |doi = 10.1371/journal.pbio.0040234|title = Prevalence and Evolution of Core Photosystem II Genes in Marine Cyanobacterial Viruses and Their Hosts|year = 2006|last1 = Sullivan|first1 = Matthew B.|last2 = Lindell|first2 = Debbie|author-link2=Debbie Lindell|last3 = Lee|first3 = Jessica A.|last4 = Thompson|first4 = Luke R.|last5 = Bielawski|first5 = Joseph P.|last6 = Chisholm|first6 = Sallie W.|journal = PLOS Biology|volume = 4|issue = 8|pages = e234|pmid = 16802857|pmc = 1484495 | doi-access=free }}{{cite journal |doi = 10.1073/pnas.1102164108|title = Phage auxiliary metabolic genes and the redirection of cyanobacterial host carbon metabolism|year = 2011|last1 = Thompson|first1 = L. R.|last2 = Zeng|first2 = Q.|last3 = Kelly|first3 = L.|last4 = Huang|first4 = K. H.|last5 = Singer|first5 = A. U.|last6 = Stubbe|first6 = J.|last7 = Chisholm|first7 = S. W.|journal = Proceedings of the National Academy of Sciences|volume = 108|issue = 39|pages = E757–E764|pmid = 21844365|pmc = 3182688|doi-access = free}}{{cite journal |doi = 10.1016/j.cub.2011.11.055|title = Marine Viruses Exploit Their Host's Two-Component Regulatory System in Response to Resource Limitation|year = 2012|last1 = Zeng|first1 = Qinglu|last2 = Chisholm|first2 = Sallie W.|journal = Current Biology|volume = 22|issue = 2|pages = 124–128|pmid = 22244998|s2cid = 7692657|doi-access = free| bibcode=2012CBio...22..124Z |hdl = 1721.1/69047|hdl-access = free}}{{cite journal |doi = 10.1038/ismej.2013.4|title = Structure and function of a cyanophage-encoded peptide deformylase|year = 2013|last1 = Frank|first1 = Jeremy A.|last2 = Lorimer|first2 = Don|last3 = Youle|first3 = Merry|last4 = Witte|first4 = Pam|last5 = Craig|first5 = Tim|last6 = Abendroth|first6 = Jan|last7 = Rohwer|first7 = Forest|last8 = Edwards|first8 = Robert A.|last9 = Segall|first9 = Anca M.|last10 = Burgin|first10 = Alex B.|journal = The ISME Journal|volume = 7|issue = 6|pages = 1150–1160|pmid = 23407310|pmc = 3660681| bibcode=2013ISMEJ...7.1150F }} Cultivation-independent metagenomic analysis of viral communities has identified additional AMGs that are involved in motility, central carbon metabolism, photosystem I, energy metabolism, iron–sulphur clusters, anti-oxidation and sulphur and nitrogen cycling.{{cite journal |doi = 10.1371/journal.pbio.0050016|title = The Sorcerer II Global Ocean Sampling Expedition: Expanding the Universe of Protein Families|year = 2007|last1 = Yooseph|first1 = Shibu|last2 = Sutton|first2 = Granger|last3 = Rusch|first3 = Douglas B.|last4 = Halpern|first4 = Aaron L.|last5 = Williamson|first5 = Shannon J.|last6 = Remington|first6 = Karin|last7 = Eisen|first7 = Jonathan A.|last8 = Heidelberg|first8 = Karla B.|last9 = Manning|first9 = Gerard|last10 = Li|first10 = Weizhong|last11 = Jaroszewski|first11 = Lukasz|last12 = Cieplak|first12 = Piotr|last13 = Miller|first13 = Christopher S.|last14 = Li|first14 = Huiying|last15 = Mashiyama|first15 = Susan T.|last16 = Joachimiak|first16 = Marcin P.|last17 = Van Belle|first17 = Christopher|last18 = Chandonia|first18 = John-Marc|last19 = Soergel|first19 = David A.|last20 = Zhai|first20 = Yufeng|last21 = Natarajan|first21 = Kannan|last22 = Lee|first22 = Shaun|last23 = Raphael|first23 = Benjamin J.|last24 = Bafna|first24 = Vineet|last25 = Friedman|first25 = Robert|last26 = Brenner|first26 = Steven E.|last27 = Godzik|first27 = Adam|last28 = Eisenberg|first28 = David|last29 = Dixon|first29 = Jack E.|last30 = Taylor|first30 = Susan S.|journal = PLOS Biology|volume = 5|issue = 3|pages = e16|pmid = 17355171|pmc = 1821046|display-authors = 1 | doi-access=free }}{{cite journal |doi = 10.1038/nature06810|title = Functional metagenomic profiling of nine biomes|year = 2008|last1 = Dinsdale|first1 = Elizabeth A.|last2 = Edwards|first2 = Robert A.|last3 = Hall|first3 = Dana|last4 = Angly|first4 = Florent|last5 = Breitbart|first5 = Mya|last6 = Brulc|first6 = Jennifer M.|last7 = Furlan|first7 = Mike|last8 = Desnues|first8 = Christelle|last9 = Haynes|first9 = Matthew|last10 = Li|first10 = Linlin|last11 = McDaniel|first11 = Lauren|last12 = Moran|first12 = Mary Ann|last13 = Nelson|first13 = Karen E.|last14 = Nilsson|first14 = Christina|last15 = Olson|first15 = Robert|last16 = Paul|first16 = John|last17 = Brito|first17 = Beltran Rodriguez|last18 = Ruan|first18 = Yijun|last19 = Swan|first19 = Brandon K.|last20 = Stevens|first20 = Rick|last21 = Valentine|first21 = David L.|last22 = Thurber|first22 = Rebecca Vega|last23 = Wegley|first23 = Linda|last24 = White|first24 = Bryan A.|last25 = Rohwer|first25 = Forest|journal = Nature|volume = 452|issue = 7187|pages = 629–632|pmid = 18337718|bibcode = 2008Natur.452..629D|s2cid = 4421951}}{{cite journal |doi = 10.1016/j.tim.2016.06.006|title = Virocell Metabolism: Metabolic Innovations During Host–Virus Interactions in the Ocean|year = 2016|last1 = Rosenwasser|first1 = Shilo|last2 = Ziv|first2 = Carmit|last3 = Creveld|first3 = Shiri Graff van|last4 = Vardi|first4 = Assaf|journal = Trends in Microbiology|volume = 24|issue = 10|pages = 821–832|pmid = 27395772}} Interestingly, a recent analysis of Pacific Ocean Virome data identified niche-specialised AMGs that contribute to depth-stratified host adaptations.{{cite journal |doi = 10.1038/ismej.2014.143|title = Depth-stratified functional and taxonomic niche specialization in the 'core' and 'flexible' Pacific Ocean Virome|year = 2015|last1 = Hurwitz|first1 = Bonnie L.|last2 = Brum|first2 = Jennifer R.|last3 = Sullivan|first3 = Matthew B.|journal = The ISME Journal|volume = 9|issue = 2|pages = 472–484|pmid = 25093636|pmc = 4303639| bibcode=2015ISMEJ...9..472H }} Given that microbes drive global biogeochemical cycles, and viruses infect a large fraction of microbes at any given time,{{cite journal |doi = 10.1128/MMBR.64.1.69-114.2000|title = Virioplankton: Viruses in Aquatic Ecosystems|year = 2000|last1 = Wommack|first1 = K. Eric|last2 = Colwell|first2 = Rita R.|journal = Microbiology and Molecular Biology Reviews|volume = 64|issue = 1|pages = 69–114|pmid = 10704475|pmc = 98987}} viral-encoded AMGs must play important roles in global biogeochemistry and microbial metabolic evolution.

Mangrove forests are the only woody halophytes that live in salt water along the world's subtropical and tropical coastlines. Mangroves are one of the most productive and ecologically important ecosystems on earth. The rates of primary production of mangroves equal those of tropical humid evergreen forests and coral reefs.{{cite journal |doi = 10.4155/cmt.12.20|title = Carbon sequestration in mangrove forests|year = 2012|last1 = Alongi|first1 = Daniel M.|journal = Carbon Management|volume = 3|issue = 3|pages = 313–322|s2cid = 153827173|doi-access = free| bibcode=2012CarM....3..313A }} As a globally relevant component of the carbon cycle, mangroves sequester approximately 24 million metric tons of carbon each year.{{cite journal |doi = 10.1007/s00114-001-0283-x|title = Relevance of mangroves for the production and deposition of organic matter along tropical continental margins|year = 2002|last1 = Jennerjahn|first1 = Tim C.|last2 = Ittekkot|first2 = Venugopalan|journal = Naturwissenschaften|volume = 89|issue = 1|pages = 23–30|pmid = 12008969|bibcode = 2002NW.....89...23J|s2cid = 33556308}} Most mangrove carbon is stored in soil and sizable belowground pools of dead roots, aiding in the conservation and recycling of nutrients beneath forests.{{cite journal |doi = 10.1007/s00468-002-0206-2|title = Nutrient partitioning and storage in arid-zone forests of the mangroves Rhizophora stylosa and Avicennia marina|year = 2003|last1 = Alongi|first1 = Daniel M.|last2 = Clough|first2 = Barry F.|last3 = Dixon|first3 = Paul|last4 = Tirendi|first4 = Frank|journal = Trees|volume = 17| issue=1 |pages = 51–60| bibcode=2003Trees..17...51A |s2cid = 23613917}} Although mangroves cover only 0.5% of the earth's coastal area, they account for 10–15% of the coastal sediment carbon storage and 10–11% of the total input of terrestrial carbon into oceans.{{cite journal |doi = 10.1146/annurev-marine-010213-135020|title = Carbon Cycling and Storage in Mangrove Forests|year = 2014|last1 = Alongi|first1 = Daniel M.|journal = Annual Review of Marine Science|volume = 6|pages = 195–219|pmid = 24405426|bibcode = 2014ARMS....6..195A|doi-access = free}} The disproportionate contribution of mangroves to carbon sequestration is now perceived as an important means to counterbalance greenhouse gas emissions.

File:Avicennia marina chloroplast genome.png{{cite journal | last1=Natarajan | first1=Purushothaman | last2=Murugesan | first2=Ashok Kumar | last3=Govindan | first3=Ganesan | last4=Gopalakrishnan | first4=Ayyaru | last5=Kumar | first5=Ravichandiran | last6=Duraisamy | first6=Purushothaman | last7=Balaji | first7=Raju | last8=Shyamli | first8=Puhan Sushree | last9=Parida | first9=Ajay K. | last10=Parani | first10=Madasamy | title=A reference-grade genome identifies salt-tolerance genes from the salt-secreting mangrove species Avicennia marina | journal=Communications Biology | publisher=Springer Science and Business Media LLC | volume=4 | issue=1 | date=8 July 2021 | page=851 | issn=2399-3642 | doi=10.1038/s42003-021-02384-8| pmid=34239036 | pmc=8266904 }} 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].]]

Despite the ecological importance of the mangrove ecosystem, knowledge of mangrove biodiversity is notably limited. Previous reports mainly investigated the biodiversity of mangrove fauna, flora, and bacterial communities.{{cite journal |doi = 10.1111/j.1574-6941.2008.00519.x|title = Exploring the diversity of bacterial communities in sediments of urban mangrove forests|year = 2008|last1 = Marcial Gomes|first1 = Newton C.|last2 = Borges|first2 = Ludmila R.|last3 = Paranhos|first3 = Rodolfo|last4 = Pinto|first4 = Fernando N.|last5 = Mendonã§a-Hagler|first5 = Leda C. S.|last6 = Smalla|first6 = Kornelia|journal = FEMS Microbiology Ecology|volume = 66|issue = 1|pages = 96–109|pmid = 18537833| bibcode=2008FEMME..66...96M | s2cid=40733636 }}{{cite journal |doi = 10.1371/journal.pone.0038600|doi-access = free|title = The Microbiome of Brazilian Mangrove Sediments as Revealed by Metagenomics|year = 2012|last1 = Andreote|first1 = Fernando Dini|last2 = Jiménez|first2 = Diego Javier|last3 = Chaves|first3 = Diego|last4 = Dias|first4 = Armando Cavalcante Franco|last5 = Luvizotto|first5 = Danice Mazzer|last6 = Dini-Andreote|first6 = Francisco|last7 = Fasanella|first7 = Cristiane Cipola|last8 = Lopez|first8 = Maryeimy Varon|last9 = Baena|first9 = Sandra|last10 = Taketani|first10 = Rodrigo Gouvêa|last11 = De Melo|first11 = Itamar Soares|journal = PLOS ONE|volume = 7|issue = 6|pages = e38600|pmid = 22737213|pmc = 3380894|bibcode = 2012PLoSO...738600A}}{{Cite book|url=https://books.google.com/books?id=EgvLwAEACAAJ&q=%22Species+diversity+in+ecological+communities:+historical+and+geographical+perspectives%22|title = Species Diversity in Ecological Communities: Historical and Geographical Perspectives|isbn = 9780226718231|last1 = Ricklefs|first1 = Robert E.|last2 = Schluter|first2 = Dolph|year = 1993| publisher=University of Chicago Press }} Particularly, little information is available about viral communities and their roles in mangrove soil ecosystems.{{cite journal |doi = 10.1016/j.tim.2017.12.004|title = The 'Neglected' Soil Virome – Potential Role and Impact|year = 2018|last1 = Pratama|first1 = Akbar Adjie|last2 = Van Elsas|first2 = Jan Dirk|journal = Trends in Microbiology|volume = 26|issue = 8|pages = 649–662|pmid = 29306554|s2cid = 25057850}}{{cite journal |doi = 10.1146/annurev-virology-101416-041639|title = Viruses in Soil Ecosystems: An Unknown Quantity within an Unexplored Territory|year = 2017|last1 = Williamson|first1 = Kurt E.|last2 = Fuhrmann|first2 = Jeffry J.|last3 = Wommack|first3 = K. Eric|last4 = Radosevich|first4 = Mark|journal = Annual Review of Virology|volume = 4|issue = 1|pages = 201–219|pmid = 28961409|doi-access = free}} In view of the importance of viruses in structuring and regulating host communities and mediating element biogeochemical cycles, exploring viral communities in mangrove ecosystems is essential. Additionally, the intermittent flooding of sea water and resulting sharp transition of mangrove environments may result in substantially different genetic and functional diversity of bacterial and viral communities in mangrove soils compared with those of other systems.{{cite journal |doi = 10.1007/s00227-006-0377-2|title = Recovery of novel bacterial diversity from mangrove sediment|year = 2007|last1 = Liang|first1 = Jun-Bin|last2 = Chen|first2 = Yue-Qin|last3 = Lan|first3 = Chong-Yu|last4 = Tam|first4 = Nora F. Y.|last5 = Zan|first5 = Qi-Jie|last6 = Huang|first6 = Li-Nan|journal = Marine Biology|volume = 150|issue = 5|pages = 739–747| bibcode=2007MarBi.150..739L |s2cid = 85384181}}

=Genome sequencing=

  • Rhizophoreae as revealed by whole-genome sequencing{{cite journal | last1=Xu | first1=Shaohua | last2=He | first2=Ziwen | last3=Zhang | first3=Zhang | last4=Guo | first4=Zixiao | last5=Guo | first5=Wuxia | last6=Lyu | first6=Haomin | last7=Li | first7=Jianfang | last8=Yang | first8=Ming | last9=Du | first9=Zhenglin | last10=Huang | first10=Yelin | last11=Zhou | first11=Renchao | last12=Zhong | first12=Cairong | last13=Boufford | first13=David E | last14=Lerdau | first14=Manuel | last15=Wu | first15=Chung-I | last16=Duke | first16=Norman C. | last17=Shi | first17=Suhua | title=The origin, diversification and adaptation of a major mangrove clade (Rhizophoreae) revealed by whole-genome sequencing | journal=National Science Review | publisher=Oxford University Press (OUP) | volume=4 | issue=5 | date=5 June 2017 | issn=2095-5138 | doi=10.1093/nsr/nwx065 | pages=721–734| pmid=31258950 | pmc=6599620 }}

{{clear}}

See also

References

{{reflist}}

Further reading

  • Saenger, Peter (2002). Mangrove Ecology, Silviculture, and Conservation. Kluwer Academic Publishers, Dordrecht. {{ISBN|1-4020-0686-1}}.
  • Thanikaimoni, Ganapathi (1986). Mangrove Palynology UNDP/UNESCO and the French Institute of Pondicherry, ISSN 0073-8336 (E).
  • Tomlinson, Philip B. (1986). The Botany of Mangroves. Cambridge University Press, Cambridge, {{ISBN|0-521-25567-8}}.
  • Teas, H. J. (1983). Biology and Ecology of Mangroves. W. Junk Publishers, The Hague. {{ISBN|90-6193-948-8}}.
  • {{cite journal |doi=10.1023/A:1011118204434 |year=2001 |last1=Plaziat |first1=Jean-Claude |journal=Wetlands Ecology and Management |volume=9 |issue=3 |pages=161–180 |last2=Cavagnetto |first2=Carla |last3=Koeniguer |first3=Jean-Claude |last4=Baltzer |first4=Frédéric |title=History and biogeography of the mangrove ecosystem, based on a critical reassessment of the paleontological record|s2cid=24980831 }}
  • {{cite journal | last1 = Jayatissa | first1 = L. P. | last2 = Dahdouh-Guebas | first2 = F. | last3 = Koedam | first3 = N. | year = 2002 | title = A review of the floral composition and distribution of mangroves in Sri Lanka | url = https://dipot.ulb.ac.be/dspace/bitstream/2013/46624/1/Jayatissaetal_2002_BotJLinnSoc.pdf| journal = Botanical Journal of the Linnean Society | volume = 138 | pages = 29–43 | doi=10.1046/j.1095-8339.2002.00002.x| doi-access = free }}
  • {{cite journal | last1 = Ellison | first1 = Aaron M. | year = 2000 | title = Mangrove Restoration: Do We Know Enough? | journal = Restoration Ecology | volume = 8 | issue = 3| pages = 219–229 | doi = 10.1046/j.1526-100x.2000.80033.x | bibcode = 2000ResEc...8..219E | s2cid = 86352384 }}
  • Agrawala, Shardul; Hagestad; Marca; Koshy, Kayathu; Ota, Tomoko; Prasad, Biman; Risbey, James; Smith, Joel; Van Aalst, Maarten. 2003. Development and Climate Change in Fiji: Focus on Coastal Mangroves. Organisation of Economic Co-operation and Development, Paris, Cedex 16, France.
  • {{cite journal | last1 = Barbier | first1 = E. B. | last2 = Sathirathai | first2 = S. | year = 2001 | title = Valuing Mangrove Conservation in Southern Thailand | journal = Contemporary Economic Policy | volume = 19 | issue = 2| pages = 109–122 | doi = 10.1111/j.1465-7287.2001.tb00054.x }}
  • {{cite journal | last1 = Bosire | first1 = J. O. | last2 = Dahdouh-Guebas | first2 = F. | last3 = Jayatissa | first3 = L. P. | last4 = Koedam | first4 = N. | last5 = Lo Seen | first5 = D. | last6 = Nitto | first6 = Di D. | year = 2005 | title = How Effective were Mangroves as a Defense Against the Recent Tsunami? | doi = 10.1016/j.cub.2005.06.008 | journal = Current Biology | volume = 15 | issue = 12| pages = R443–R447 | pmid = 15964259 | s2cid = 8772526 | url = http://agritrop.cirad.fr/529549/ | doi-access = free }}
  • {{cite journal | last1 = Bowen | first1 = Jennifer L. | last2 = Valiela | first2 = Ivan | last3 = York | first3 = Joanna K. | year = 2001 | title = Mangrove Forests: One of the World's Threatened Major Tropical Environments | journal = BioScience | volume = 51 | issue = 10| pages = 807–815 | doi = 10.1641/0006-3568(2001)051[0807:mfootw]2.0.co;2 | doi-access = free }}
  • {{cite journal | last1 = Jin-Eong | first1 = Ong | year = 2004 | title = The Ecology of Mangrove Conservation and Management | journal = Hydrobiologia | volume = 295 | issue = 1–3| pages = 343–351 | doi = 10.1007/BF00029141 | s2cid = 26686381 }}
  • Glenn, C. R. 2006. [http://earthsendangered.com "Earth's Endangered Creatures"]
  • {{cite journal | last1 = Lewis | first1 = Roy R. III | year = 2004 | title = Ecological Engineering for Successful Management and Restoration of Mangrove Forest | journal = Ecological Engineering | volume = 24 | issue = 4| pages = 403–418 | doi = 10.1016/j.ecoleng.2004.10.003 }}
  • {{cite journal |last1=Kuenzer |first1=C. |last2=Bluemel |first2=A. |last3=Gebhardt |first3=S. |last4=Vo Quoc |first4=T. |name-list-style=amp |last5=Dech |first5=S. |date=2011 |title=Remote Sensing of Mangrove Ecosystems: A Review |journal=Remote Sensing |volume=3 |issue=5 |pages=878–928 |doi=10.3390/rs3050878|bibcode=2011RemS....3..878K |doi-access=free }}
  • {{cite journal | last1 = Lucien-Brun | first1 = H | year = 1997 | title = Evolution of world shrimp production: Fisheries and aquaculture | journal = World Aquaculture | volume = 28 | pages = 21–33 }}
  • Twilley, R. R., V. H. Rivera-Monroy, E. Medina, A. Nyman, J. Foret, T. Mallach, and L. Botero. 2000. Patterns of forest development in mangroves along the San Juan River estuary, Venezuela. Forest Ecology and Management
  • {{cite journal | last1 = Murray | first1 = M. R. | last2 = Zisman | first2 = S. A. | last3 = Furley | first3 = P. A. | last4 = Munro | first4 = D. M. | last5 = Gibson | first5 = J. | last6 = Ratter | first6 = J. | last7 = Bridgewater | first7 = S. | last8 = Mity | first8 = C. D. | last9 = Place | first9 = C. J. | year = 2003 | title = The Mangroves of Belize: Part 1. Distribution, Composition and Classification | journal = Forest Ecology and Management | volume = 174 | issue = 1–3| pages = 265–279 | doi=10.1016/s0378-1127(02)00036-1| bibcode = 2003ForEM.174..265M }}
  • {{cite journal |last1=Vo Quoc |first1=T. |last2=Kuenzer |first2=C. |last3=Vo Quang |first3=M. |last4=Moder |first4=F. |name-list-style=amp |last5=Oppelt |first5=N. |date=December 2012 |title=Review of Valuation Methods for Mangrove Ecosystem Services |journal=Ecological Indicators |volume=23 |pages=431–446 |doi=10.1016/j.ecolind.2012.04.022|bibcode=2012EcInd..23..431V }}
  • Spalding, Mark; Kainuma, Mami and Collins, Lorna (2010) World Atlas of Mangroves Earthscan, London, {{ISBN|978-1-84407-657-4}}; 60 maps showing worldwide mangrove distribution
  • Warne, Kennedy (2013) Let them eat shrimp: the tragic disappearance of the rainforests of the sea. Island Press, 2012, {{ISBN|978-1597263344}}
  • {{cite journal | last1 = Massó | last2 = Alemán | first2 = S. | last3 = Bourgeois | first3 = C. | last4 = Appeltans | first4 = W. | last5 = Vanhoorne | first5 = B. | last6 = De Hauwere | first6 = N. | last7 = Stoffelen | first7 = P. | last8 = Heaghebaert | first8 = A. | last9 = Dahdouh-Guebas | first9 = F. | year = 2010 | title = The 'Mangrove Reference Database and Herbarium' | url = http://www.vliz.be/imisdocs/publications/236913.pdf| journal = Plant Ecology and Evolution | volume = 143 | issue = 2| pages = 225–232 | doi = 10.5091/plecevo.2010.439}}
  • {{cite journal |last1=Vo Quoc |first1=T. |last2=Oppelt |first2=N. |last3=Leinenkugel |first3=P. |name-list-style=amp |last4=Kuenzer |first4=C. |date=2013 |title=Remote Sensing in Mapping Mangrove Ecosystems – An Object-Based Approach |journal=Remote Sensing |volume=5 |issue=1 |pages=183–201 |doi=10.3390/rs5010183|bibcode=2013RemS....5..183V |doi-access=free }}