Palm snorkelling: leaf bases as aeration structures in the mangrove palm (Nypa fruticans) G. Chomicki, Luc Bidel, William. J. Baker, Christian Jay-Allemand

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G. Chomicki, Luc Bidel, William. J. Baker, Christian Jay-Allemand. Palm snorkelling: leaf bases as aeration structures in the mangrove palm (Nypa fruticans). Botanical Journal of the Linnean Society, Linnean Society of London, 2014, 174 (2), pp.257-270. ￿10.1111/boj.12133￿. ￿hal-01189916￿

HAL Id: hal-01189916 https://hal.archives-ouvertes.fr/hal-01189916 Submitted on 27 May 2020

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Version preprint 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm bs_bs_query bs_bs_query Botanical Journal of theLinnean Society, 174(2), 257-270. DOI :10.1111/boj.12133 2 1 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). farto tutrs hssuypoie tutrleiec niaigthat persistence the indicating ensuring evidence abscission, following structural years aeren- provides tissues. 4 living study to schizo-lysigenous to up This of transport for structures. oxygen live consists longitudinal can aeration cortex facilitating bases Expansigenous of branching and leaf the root that base. loss and two suggests oxygen lenticels leaf first estimation radial Simultaneously, exodermis, the The Lifespan limiting base. and base. covers lacuna, leaf leaf wide lenticels rhizodermis the with the mature in subero-lignified chyma occurs of with a junction network junction stem present a the of orders towards completed, zone porosity is increasing the abscission with at aerenchyma when abscised that, is so rachis develop the senescence, Following 03,Germany 80638, HITA JAY-ALLEMAND CHRISTIAN 2 1 CHOMICKI GUILLAUME in fruticans) structures (Nypa aeration palm as mangrove bases the leaf snorkelling: Palm 2013, Society, Linnean the of Journal Botanical 03TeLnenSceyo London, of Society Linnean The 2013 © [email protected] [email protected]; E-mail: author. *Corresponding pneumato- include These adaptations [e.g. 1995). phores root Tomlinson, aerial 1955; typically Scholander, Van & (Scholander, systems Dam root morphologi- their of remarkable adaptations the cal to is environments attributed reduced, and classically highly waterlogged, and in anoxic species therefore mangrove of success The London, of Society Linnean The KEYWORDS: 2013 ADDITIONAL © 2013, Society, habitat. Linnean mangal the the in of structure air-supplying of type but, roots, the to of oxygen supply adaptations to pneumatophores, the as such structures, specialized in evolved 2013 have species October Mangrove 29 publication for accepted 2013; October 28 revised 2013; July 31 Received 4 3 ame–rci bcsin–ro xgnspl ciolsgn tannins. – schizo-lysigeny – supply oxygen root – abscission rachis – Palmae – ihbace,sc si h e age(Rhizophora mangle from red the ectopically mangle in emerge as such can branches, that high roots aerial titious ytmtcBtn n yooy eateto ilg,Uiest fMnc LU,Munich France (LMU), Montpellier, Munich AGAP, of UMR University Montpellier, INRA Biology, of Department Mycology, and Botany Systematic nvriyo otele ,URDAE lc uèeBtiln -405 otele,France Montpellier, 095, F-34 Bataillon, Eugène Place DIADE, UK UMR 3AB, 2, TW9 Montpellier Surrey, of Richmond, University Kew, Gardens, Botanic Royal h nymnrv am osc tutr a enrpre.Ti td ie odetermine to aimed study This reported. been has structure such no palm, mangrove only the fruticans, Nypa .,o omacutra h aeo h trunk, the of base the at cluster a form or L.), vcni marina Avicennia INTRODUCTION .fruticans N. Comment citer cedocument: 000–000. 00, Frk)Veh] adven- Vierh.], (Forsk.) eecya–Aeaee–epniey–hprrpi etcl–la lifespan leaf – lenticel hypertrophic – expansigeny – Arecaceae – aerenchyma otemna aia ihseilrfrnet h i-upyn structure. air-supplying the to reference special with habitat mangal the to FLS

2013, Society, Linnean the of Journal Botanical 4 1 ,LCP .BIDEL R. P. LUC *, ••.Wt figures 8 With ••–••. ••, omto farnhm ihitronce air interconnected with that aerenchyma the trait is of anoxia anatomical soil and formation key inundation to the tolerance enables systems, root grove 1995). Tomlinson, 1982; to (Scholander Saenger, environment metabolism transport anaerobic such aerobic an oxygen that in ensuring hence assumed enable tissues, been systems drowned long root man- has In adaptive it 1995). species, (Tomlinson, grove entry whilst water transfer gas preventing enable high which lenticels, possess of structures densities these of All 1995). Tomlinson, of those of roots, as such littoralis roots, [e.g. buttress in roots Lamk.], as such knee roots, Griff, stilt as known eodtemrhlgclaattoso man- of adaptations morphological the Beyond 2 ILA .BAKER J. WILLIAM , io,o iuu utess ocle plank so-called buttresses, sinuous or Aiton, yoapsgranatum Xylocarpus .fruticans N. ••–•• ••, rgir gymnorrhiza Bruguiera il Segr 1982; (Saenger, Ridl. a vle unique a evolved has FLS hzpoastylosa Rhizophora oaia Journal Botanical 3 and ,1955; al., et Heritiera (L.) 1 Version preprint 56 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm Botanical Journal of theLinnean Society, 174(2), 257-270. DOI :10.1111/boj.12133 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). bs_bs_query bs_bs_query 4 3 oly 00 e,20;Hre,20;Dransfield 2006; Harley, Gomez-Navarro 2008; 2001; al., et Gee, 1964; 2000; (Tralau, (Maastrichtian) Morley, Cretaceous upper the 2013). Couvreur, & early (Baker an Mya 93.5 as phylogeny palm arose the of that (Asmussen base genus the near monotypic lineage diverging a and is Australia (Dransfield Steck to Islands Asia in Solomon South-East Lanka the Sri through and west India from the distributed is and palm, (Scholander 2001). at peaking depend- tides concentration is oxygen low system with the root tides, the species, upon ent of mangrove oxygenation in of pattern Moreover, 2005). Suzuki, eln ntedsrbto of of late dramatic records distribution Gomez-Navarro the a 2006; the in Harley, with in climate coincided drier decline and Miocene a environments towards Eocene–early waterlogged changes in extinct that found pal- that also Recent indicate were Africa. range, and data modern Europe its aeobotanical Americas, the outside in far e.g. from reported rences species, mangrove several for detail including in described and separa- division or collapse cell death, cell (Seago by enlarge- by tion spaces the not but involves intercellular expansion, it of that in ment layers processes two from cell different these spaces somewhat is intercellular adjacent Expansigeny of lacunae. joining into of the in separation resulting aer- occurs, schizogenous from cell in whereas, arise enchyma, collapse, lacunae and air lysis cell the aerenchyma, lysigeny, (Seago development: lysigenous expansigeny of and mode the schizogeny catego- to main Armstrong, three according oxygen 1920; into ries of fall (Arber, types rate tissues Aerenchyma the 1979). plant increase in to air diffusion of pivotal formation the are and channels porosity Consistently, tissue in plant. increase the an throughout systems channel 2 97 and 1977) tmta rnhsi ihtmu anr which, manner, dichotomous ortho- a in an branches eudicot have that with stem which system, contrast shoot shrubs, tropic and In trees 1). (Fig. mangrove plant mangrove in and adaptation genus recent the a for the not synapomorphic to is adaptation habitat, the the mangal therefore that and by suggests habit, biota This mangrove mangrove the 2007). of (Gee, marker a palaeontologists as used are ments Nypa fruticans Nypa eecyasrcueaddvlpethv been have development and structure Aerenchyma yafruticans Nypa .CHOMICKI G. a netniefsi eoddtn akto back dating record fossil extensive an has hzpoamangle Rhizophora Nypa ,20;Baker 2006; al., et ,2005). al., et vcni marina Avicennia ope ihsappalaeoenviron- swamp with coupled um.i h nyetn mangrove extant only the is Wurmb. osse nuuulhbtfra for habit unusual an possesses TAL. ET Comment citer cedocument: .fruticans N. 03TeLnenSceyo London, of Society Linnean The 2013 © ,15;Allaway 1955; al., et .fruticans. N. ,20) ihoccur- with 2009), al., et ,20) ae at dated 2009), al., et Gl n Tomlinson, and (Gill ,20) Ancient 2009). al., et

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Royal number the (accession at UK Kew, collected root Gardens, was Additional material 82473B). leaf planted number and was (accession specimen 1982 The in USA. FL, Miami, Center, specimen mature a from of collected was material Plant euto h lgorpchbtof habit plagiotropic As the termi- 1C). of Fig. that result 1971, a head (Tomlinson, maturation, fruiting inflorescence the the seed nates from following shed being of germinate before adaptation which remarkable lings One (Tomlinson, 1). Nypa colonization Fig. vegetative 1995; 1971, allows turn, in tmi fe mesd h evsbigteonly 1). 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(Tomlinson, wet pneumatophores in mangal mangrove as living other associates species palm many some like including species, roots breathing produce 2013, Society, Linnean the of Journal Botanical nti td,w xlr h oeo the of role the explore we study, this In .fruticans N. olwadsiciedvlpetlpt,whereby path, developmental distinctive a follow otemna aia stevvpru seed- viviparous the is habitat mangal the to A NATOMY AEILADMETHODS AND MATERIAL .fruticans N. rwn tteMngmr Botanical Montgomery the at growing , P TIIGADMICROSCOPY AND STAINING ATMATERIAL LANT Nypa a vle nair-supplying an evolved has evspa h oeo a of role the play leaves Nypa Nypa n ie,its tides, and Nypa ••–•• ••, osnot does .fruti- N. efin leaf 110 109 108 107 106 105 104 103 102 101 100 99 98 97 96 95 94 93 92 91 90 89 88 87 86 85 84 83 82 81 80 79 78 77 76 75 74 73 72 71 70 69 68 67 66 65 64 63 62 61 60 59 58 57 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Version preprint 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm bs_bs_query 14 bs_bs_query bs_bs_query Botanical Journal of theLinnean Society, 174(2), 257-270. DOI :10.1111/boj.12133 5 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). Colour online, B&W in print efwt h w nlsn roe.Saebr:B 5c;C ,1 cm. 10 D, C, cm; 15 B, bars: Scale grooves. enclosing two the with leaf of of cences) morphology general and Habit 03TeLnenSceyo London, of Society Linnean The 2013 Rechnitz, © & (Sheahan and 1992). quercetin respectively) (typically ester kaempferol, flavonoids with is 2-aminoethyl fluorescence associated distribution yellow–green or flavonoid acid yellow–gold tissues; seek in to was diphenylboric slides nm) (DPBA)-stained 450–480 on spectrum, used (excitation (Liu filter walls B2-A cell autofluorescence other blue Under all and depicts tissues. 380– autofluorescence in lignin attributed lignin is to spectrum, fluorescence track blue–white to bright filter, (4′,6-diamidino-2- used this (excitation was DAPI nm) Nikon 420 filter A 1. phenylindole) Ds-Fi camera 1. Figure ,Iflrsec.D on ihtmzn tmsoigtetosse hosadtesubtending the and shoots sister two the showing stem dichotomizing Young D, Inflorescence. C, fruticans . N. yafruticans Nypa Comment citer cedocument: ai,seie rwn tteMngmr oaia etr L S,patdi 92 A, 1982. in planted USA, FL, Center, Botanical Montgomery the at growing specimen habit, ,Yugiflrsec hwn h loati ai ailr inflores- (axillary habit pleonanthic the showing inflorescence Young B, fruticans . N. ,19) Nikon A 1994). al., et

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DOI :10.1111/boj.12133 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). Colour online, B&W in print 03TeLnenSceyo London, of Society Linnean The 2013 in © lifespan base leaf The time. long Nypa a senesce, for not persist do but bases leaf abscission, rachis Following 60 D, cm; E. 1.2 in B, shown a cm; zone 10 Deg represent A, the panel bars: in Top abscission Scale during zone. agglomeration. hydrolysis abscission cellulose wall the Cell and J, in wall non-lignified I, Young cells cell G, plane. degraded E). divided division 200 in the the the Recently point zone to in show Arrowheads H, Deg parallel image. section Arrowheads (4′,6-diamidino-2-phenylindole) the walls E). 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2013, Society, Linnean the of Journal Botanical yafruticans Nypa ;E ,120 F, E, μm; ot.Bakarw indicate arrows Black roots. ;G ,70 H, G, μm; ••–•• ••, ;I–M, μm; Version preprint 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm bs_bs_query 14 bs_bs_query Botanical Journal of theLinnean Society, 174(2), 257-270. DOI :10.1111/boj.12133 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). Colour online, B&W in print 03TeLnenSceyo London, of Society Linnean The 2013 © species, other in palm abscission the the in role including as by key a such play degrade to been known long enzymes, to have which wall-degrading easy polygalacturonase, and cellulase cell abscission, of of activation time the abscission at becomes, the tissue of ground cellulose-rich determinant This crucial mechanism. a zone is abscission 2E–G) the (Fig. in cells non-lignified thick-walled leaf. whole the not and concerns rachis abscission the because only categories these the into fall Clearly, by exhibited 1990). pattern Tomlinson, abscission 1915; Floresta, Schoute, (La structure 1904; sheath the to related as palm trunks, non-self-cleaning recognized versus palm self-cleaning however, with of ecologically was, early area dichotomy and major neglected A a biology. developmentally is abscission, phenomenon, a leaf important being that despite emphasized (1990) Tomlinson the of position the illustrates left top the at scheme The C. (basal-most). and oldest B the A, C bases and leaf (distal-most) youngest the being 8. Figure n srdcdt ml neclua pcs(Fig. spaces intercellular small to reduced non-schizogenous is and non-lysigenous and is (AX3) order E F). 7E, h rsneo o-infidfirsado larger of and fibres non-lignified of presence The ncnrs,arnhm ntetidadls root last and third the in aerenchyma contrast, In annacmlto ntela aeof base leaf the in accumulation Tannin T EASISO YEIN TYPE ABSCISSION HE DISCUSSION lesguineensis Elaeis Comment citer cedocument: .fruticans N.

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DOI :10.1111/boj.12133 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). lopeeti h tmo amseissc as such species palm of porosity stem the increasing in is of aerenchyma present of type also way This junction. stem efficient of the towards bases an leaf in provides present root aerenchyma in sigenous stage (1996). Seubert early by noticed an also relation at this was separation; present cell expansigeny), facilitating be development, radial must (i.e. which of cells spaces parenchymatous intercellular by the promoted particularly be 1996). appeared to Schizogeny family. the Seubert, in aerenchyma this of for 1959; type evidence unequivocal first Udwadia, the is this & Therefore, evidence Mahabale microscopic provided (e.g. has none roots, schizogeny palm demonstrate in to Although claimed roots. have palm studies in other schizogeny of speculated & existence (1990) the lysig- Horn on Tomlinson of Tomlinson, although is 2011), 1990; species, Fisher, (Tomlinson, palm many origin of enous roots the in which occurs development, aerenchyma that argued been has nterosof roots the aerenchyma in lysigenous third-order described (1904) the Drabble and in root. roots; aerenchyma second-order expansigenous and radial first- schizo-lysigenous the rachis; in the of aerenchyma part proximal the and base leaf the in in aerenchyma identified expansigenous be honeycomb can aerenchyma Seago of of types taxonomy the Following long the in a of role lifespan a probably plays Thus, probably is accumulation pathogens. tannin accumulation such against tannin Therefore, mechanism protective of pathogens. semi- to process and sensitive this them heterotrophic makes of The sion bases leaf of 8). condition (Fig. leaf immersed of aging marker a base is accumulation most gradual subsequent the therefore (Satar and most tannin-rich the corrosion, are against stem inflorescence proficient and base leaf the of that solutions base. tannin of leaf potential the in is role tissue protective ground a base generalized with leaf The consistent in 1989). tannins Wada, of & accumulation with Mackie, (Kimura association ions by & conditions ferric soil Smith acidic and 2003; pathogens reduced Mackie, fungal Ishida & 2004; and Imlay bacterial (Smith, protective play against to shown functions of been have bark tannins oxidants, and Tomlinson, leaves has 1989; Hernes Wada, 1995; roots, tannins & (Kimura in of tannins. species mangrove reported role likely most widely protective are been and and occurrence red, The dark and soluble 12 h eeomna rdeto oecm expan- honeycomb of gradient developmental The eetsuyeautn h orso inhibition corrosion the evaluating study recent A .CHOMICKI G. 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Beckett, previous Saenger, & with greatly Armstrong (Armstrong, consistent certainly to entry, helps (AX3) phytotoxin minimize probably that roots conditions This oxidizing microaerobic loss. create fine oxygen radial the enhances surface on hydrophobic enhancing of ratio, layers absence volume The to uptake. root also surface nutrient flooded certainly large roots the a fine of of produces number demand large This oxygen system. the reduce in may system root less 1:2:10 the much of consume architecture roots (Bidel roots fine major of than that oxygen ratio shown 10 been : in 1 has roots This fine 7B) over (Fig. large orders branching three of stem. stem the the of that practical known junction for is here the studied it not to reasons, was down stem the oxygen Although conduction the a allows expansig- of base leaf the the in by aerenchyma provided enous lenticels, lacuna from and interconnected entry the mangrove air Following in 1979). tissues Armstrong, (Scholander into plants woody entry non-mangrove air enhance known been to long have which into lenticels, hypertrophic entry by Oxygen pathway. of base air leaf aeration the the an for as account base nary leaf the of in role structure the to addition In hssuypoie tutrleiec o new a for species. mangrove evidence a of structural structure aeration provides study This A 2013, Society, Linnean the of Journal Botanical unicto ftebaciglvlo ahroot each of level branching the of Quantification RPTWYFO H EFBS OTEROOTS THE TO BASE LEAF THE FROM PATHWAY IR bnac ftetrero rnhn orders, branching root three the of abundance osoe regia Roystonea hssuypoie prelimi- a provides study this fruticans, N. 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(F.Muell.) ,20) hs the Thus, 2000). al., et 1:2:10 ihthe with Nypa, Nypa ,2011). al., et Nypa ,1955; al., et ai fthe of ratio ••–•• ••, salso is leaves tal., et 111 110 109 108 107 106 105 104 103 102 101 100 99 98 97 96 95 94 93 92 91 90 89 88 87 86 85 84 83 82 81 80 79 78 77 76 75 74 73 72 71 70 69 68 67 66 65 64 63 62 61 60 59 58 57 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Version preprint 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm Botanical Journal of theLinnean Society, 174(2), 257-270. DOI :10.1111/boj.12133 snorkelling: leafbases asaerationstructures inthe mangrove palm (Nypafruticans). lopoie h potnt oivsiaeit. investigate and collect permission to to for help opportunity acknowledged technical is but the Griffith pro- study, Patrick only this provided not in who investigated also Tomlinson, hypothesis B. the P. posed expresses to author gratitude first The his critical manuscript. for the Aroles of Jeremy reading and Rudall Paula thank We of physiology the Further investigate in aeration time. oxygenation to root long the needed a are that for studies pro- functional ensuring by remains tannins, aided system perhaps for from abscission, alive after remain tection bases years 4 Leaf roots. to from the up to occurs lenticels base that leaf network place. hypertrophied whilst the aerenchyma in base the These to leaf base senesces. connect the leaf rachis on the develops the leaving only network which shed, lenticel by A is process rachis abscission the unique a undergo 03TeLnenSceyo London, of Society Linnean The 2013 © 1979. W. Armstrong 1992. PM. Beckett W, Armstrong J, Armstrong 1920. A. Arber MC, Ricketts LM, Hollington M, Curran WG, Allaway 1982. FT. Addicott ae J overT.2013. TL. Couvreur WJ, Baker AS, Barfod V, Deickman J, Dransfield CB, Asmussen oddct hspprt .B olno n his in and and, Tomlinson plants B. of are structure P. palms. the particular, to reviewers wish in paper We career anonymous remarkable manuscript. this the Two dedicate on to 2009. comments for in thanked G.C. scholarship a for to thanked is (NTBG) Gardens Tropical on USA, data phenological fruticans FL, providing N. for Miami, Noblick (MBC), Larry and Center Botanical gomery osehnerioearto n hzshr oxidation. pressure rhizosphere Phytologist and New humidity-induced aeration rhizome and enhance flows venturi- australis: mites Press. University Cambridge Cambridge: mangrove. grey the Management and Duke, Ecology NC lands ex Moldenke (Walp.) of 2001. NJ. Skelton Press. California iesfiaino am hd ih nteeouinof evolution the on geography biogeography. light Historical I. sheds lineages. tropical palms of diversification 151: plastid from evidence phylogeny. (Arecaceae): DNA family palm the 2006. of cation WJ. Baker J-C, Pintaud Research Botanical vcni marina Avicennia 15–38. 40: rwn tMC h ainlBotanical National The MBC. at growing ACKNOWLEDGEMENTS ae lns td fautcangiosperms. aquatic of study a plants: Water 274–285. 120: oaia ora fteLnenSociety Linnean the of Journal Botanical REFERENCES ekly A nvriyof University LA: Berkeley, Abscission. a pc n xgnecag nroots in exchange oxygen and space Gas 7: eaini ihrplants. higher in Aeration .fruticans. N. 197–207. 275–332. Comment citer cedocument: Frs. ir.var. Vierh. (Forssk.) Nypa 9: aeila h Mont- the at material lblboegah and biogeography Global e ufml classifi- subfamily new A 221–228.

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Tropical 2 Trans- 1086– diffu- Cel- 13 6: 9: bs_bs_query 9 113 112 111 110 109 108 107 106 105 104 103 102 101 100 99 98 97 96 95 94 93 92 91 90 89 88 87 86 85 84 83 82 81 80 79 78 77 76 75 74 73 72 71 70 69 68 67 66 65 64 63 62 61 60 59 58 57 56 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Version preprint 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm Botanical Journal of theLinnean Society, 174(2), 257-270. 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Basin, Maracaibo the in changes of roots (aeren- adventitious gas-space in and formation production chyma) ethylene of Stimulation n ooiyi oto agoeplant, mangrove a of (Lythraceae). root in porosity and 55–67. agoeeoytm nAsrla tutr,function structure, Australia: in ecosystems Mangrove .CHOMICKI G. 46: orso niiino lmnu yuignipah using by aluminium of inhibition Corrosion 5: inaelegans Zinnia 194–199. 3896–3902. and oyaatrns from polygalacturonase A hsou vulgaris Phaseolus ora fPatResearch Plant of Journal Middle 35: .anglica. S. eiwo aebtn n Palynology and Paleobotany of Review 53: opooia,aaoia n reproduc- and anatomical Morphological, ln Physiology Plant rgnadeouino rpclrain tropical of evolution and Origin yafruticans. Nypa otiuin laBooi Vegetale Biologia alla Contribuzione 101–108. Comment citer cedocument: 732–737. TAL. ET u ecnsodlacdt delle caduta della meccanismo Sul oeemnrvsadvegetation and mangroves Eocene tmtissues. stem 03TeLnenSceyo London, of Society Linnean The 2013 © ann nmnrv reroots tree mangrove in Tannins qai Botany Aquatic eecyadvlpetand development Aerenchyma ytei fcluaeduring cellulase of Synthesis rceig fteNational the of Proceedings Eocene efexplants. leaf Planta eorpi td of study demographic A cetfi eerhand Research Scientific tde npls part palms: on Studies eecyaformation Aerenchyma 53:

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Cambridge: Series. Biology Tropical Press. Clarendron Oxford: t ihtmu rnhn ntenp palm. nypa the in Botany branching dichotomous its vnk eesaskdmesHandlingar Vetenskapsakademiens Svenska ge ofloigadriopeeoiaini man- in oxidation rhizosphere and seedlings. flooding grove to egies tissue. plant of 81–90. samples small very in 1958–1967. (HCl) acid hydrochloric in media. solution extract fruticans) (Nypa xod lrno Press. Clarendon Oxford: Palmae. gastro- 1104–1115. rat the tract. in activity intestinal metabolic and diversity bacterial tannins. condensed Microbiology Environmental of effects inhibitory allows come response stress oxidative in 597–619. Phytelephantoideae. Nypoideae, aerenchyma. to respect with Botany of plants Annals flowering wetland in 2005. DE. Enstone O, Buitenzorg palmiers. quelques de mangroves. of roots Botany the in exchange and species mangrove from tannins Condensed 313. Molecules rbdpi thaliana. Arabidopsis hzpoamangle Rhizophora nentoa ora fEetohmclScience Electrochemical of Journal International 35: 42: 15: 14: 865–879. 92–98. 420–431. 57–82. otaaoyo am I.Ceroxyloideae, III. palms of anatomy Root h genus The ple niomna Microbiology Environmental Applied utainJunlo Botany of Journal Australian u afisr éin el an foliaire gaine la de médiane fissure la Sur 96: h ho pxof apex shoot The h oayo agoe.Cambridge mangroves. of botany The 565–579. ntm ftemnctldn.II. monocotyledons. the of Anatomy h tutrlbooyo palms . of biology structural The eeaiaino h otcortex root the of re-examination A nae uJri oaiu de Botanique Jardin du Annales BioTechniques n hi nixdn activity. antioxidant their and feto odne ann on tannins condensed of Effect 69: Nypa ntmclaatv strat- adaptive Anatomical 3406–3411. lvni-pcfi staining Flavonoid-specific edsRepertorium Feddes shrci coli Escherichia esrmn fporosity of Measurement van mrcnJunlof Journal American ln n Soil and Plant yafruticans Nypa 13: Wurmb. adlacandel Kandelia h ntm of anatomy The 880–883. nraigthe Increasing 5: ple and Applied 5–29. ••–•• ••, nasof Annals 44: Kungliga oover- to 297– 107: 253: Gas and 70: 7: bs_bs_query 12 104 103 102 101 100 99 98 97 96 95 94 93 92 91 90 89 88 87 86 85 84 83 82 81 80 79 78 77 76 75 74 73 72 71 70 69 68 67 66 65 64 63 62 61 60 59 58 57 56 55 54 53 bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query bs_bs_query Version preprint Chomicki, G.(Auteurde correspondance), Bidel,L., Baker,W.J., Jay-Allemand, C.(2014).Palm Botanical Journal of theLinnean Society, 174(2), 257-270. 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