Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. P I Bee ta. 09 n P V(yge l,21) upr h xaso ftetraditional the of expansion the support classification, 2016), Group’s al., Phylogeny et Angiosperm (Byng the IV studies, APG such and on 2009) al., Based et 2015). Walker (Bremer 2019; al., al., III et et Yang (Leebens-Mack APG sequencing 2018; transcriptome (Brockington, relationships al., and 2009), Sch¨aferhoff, sequencing Borsch, 2011) M¨uller, 1992; phylogenetic et gene & al., Manhart, their targeted et & (Arakaki 1994), Wilson, and sequencing Rettig, (Rodman, 2015) plastome 2011; morphology Soltis, approximately al., by & and time et Soltis families long Glover, Hernandez-Ledesma 39 Walker, a salt about over 2009; with studied whereas al., plants been flowering salt, et have of secrete (Bremer lineages major to species the glands 12,500 Polygonaceae. of related one utilise is the family Caryophyllales of The the members discuss salt-tolerant within and in Plumbaginaceae species secretory Later, absent the evolved salt-tolerant (3) in are recently tissues. structures all glands most ordinary salt-secreting that the of the are observation distribution of trichomes the only our cells glandular examine conditions the The we primitive among (4) Here the scattered role developed. structures.” In were cavities this (2) the cells and fulfil are organs. such During ducts plant to of “(1) salt secretory inside evolved groups concluded variety secrete developed having or (1988) a first that Fahn structures idioblasts tissues 1988). secrete secretory of Glands secretory Oross, diversity L¨uttge, to 1971). evolution & a Faraday of evolved 1988; with Thomson, course have (Fahn, plants 2017; salts flowering tissues Larkin, of and to specialised (Dassanayake families mucilages and the to plants across nectar found in from phenomenon substances common of a is Exudation conclude the We group sister leaves. its INTRODUCTION the and I with Polygonaceae concentrations the sister between Ca2+ the split regulate of the to members after or have place salt-tolerant might Cl- other took in glands and/or glands and whether absent salt Na+ SO42) consider however, with of and Plumbaginaceae. HCO3-, are, lineages toxicity families Cl-, of glands three the establishment the Mg2+, Salt avoid The the in Ca2+, to that glands Frankeniaceae. Zn). means K+, salt and and the a and (Na+, Tamaricaceae of Pb as ions Limoniastum related structure Ni, arisen the of closely Aegialitis, Mn, describe range the of We Fe, a in Polygonaceae. each Cu, secrete common family in Cr, also can members two Cd, are families halophytic (As, Limonium; glands three the elements saline salt genus All the and and of the Plumbago. arid glands and species in of salt Goniolimon halophytic range are have Ceratostigma, wide 45 family Armeria, a the Psylliostachys, least to in of at adapted each species species, one for and salt-tolerant known Myriolimon, its well Of family a is habitats. Caryophyllales) (non-core Plumbaginaceae The Abstract 2020 5, May 4 3 2 1 Flowers Caperta Ana and tolerance evolution stress origin, in roles Plumbaginaceae: the in structures Secretory nvriyo Sussex of University Almeria of University Tecnologias e Humanidades Lus´ofona de Universidade Lisboa de Universidade 4 1 n R´ois Ana , 2 eeoaTeixeira Generosa , 1 1 er Garcia-Caparros Pedro , 3 n i J. Tim and , Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. ihntePubgnca,mclg lns(e iue2 eedsrbd ntenntet etr,in century, nineteenth 1979). (Fahn, the cells in elongated described, radiate were several 2) supporting Figure (biseriate), (see side glands by mucilage side Plumbaginaceae, cells, are of the of examples variety Krause, Within mucilage; rows a secrete & White, two that Knox for least & (Galloway, 1979) of Fahn at ways organs Neugebauer by stipules various colleters George, ground young in (called in (Brown, below trichomes microenvironment seen have clades plant and species influence angiosperm plant above Some to all both 2020). potential in from the found with produced are 2017), be structures can ecosystems secretory plants gypsum purposes; from and secretions inland-saline The and coastal GLANDS SECRETING in MUCILAGE diversified 2 1993). have of Kubitzki, that 2015; species species al., four et halophytic (Hernandez-Ledesma the 45 of of species least one 16 at and the of 1986c) of species one Thomson, two with the together are , as 2015) al., of of et of species species species Both 2002; seven 22 halophyte. Heumann, the the a 1980; of Smith, of one & one Just Cole and Mullen, 1995). (Farago, al., see soils et heavy-metal-rich eHALOPH; Neumann on in and (K¨ohl, (recorded 1997) sandy species on genus salt-tolerant this the with of by within evolved genera and accepted species having nine been Both species have are 1). names 1500 there Table whose Howe- Plumbaginaceae, than plants tolerance. the fewer of Within of species with levels 351,000 rare approximately higher is the discriminate (http://www.theplantlist.org/). of NaCl taxonomists to 1% mM than order sepa- 80 halophytes less in that represent tolerate that (2008) mM, line so Colmer to dividing 200 (https://www.sussex.ac.uk/affiliates/halophytes/index.php?content=plantStats) & ability the ability of of Flowers as the glycophytes. matter concentration, mM Flowers, even called a salt 80 & ver, commonly been Huchzermeyer around higher species, has 2008; of a salt-sensitive salt’ Colmer, concentration used more & of salt medi- Flowers from concentration a the 2002; halophytes ‘significant selecting Breckle, in rates a (1989) salt example, Aronson to soluble for with amounts of (see, 2013) what concentrations years Quite significant the halophytes grow. of over are presence debate they that (Table the which family tolerance in a in salt survive within um display species that 5% of plants whole, proportion are a the Halophytes as on 2016). family Flowers, based 1993; list the & (Kubitzki, a Aronson in habitats in Al-Azzawi, species fourth (Santos, coastal 940 family nearly in the the Sanmart´ın, ranking often mem- Kazempour,-Osaloo, Of Moharrek, 1), its and 2019). 2017; Plumbaginoideae, Feliner, environments al., Nieto et the saline Malekmohammadi & Within and 2015; Oceania). al., arid and et in Hernandez-Ledesma Akhani, Asia occur Malekmohammadi, in 1): alt- predominantly 2005; (Table species al., regions, bers mangrove 2019) (Lled´o et Irano-Turian (two Hemisphere al., and Southern distribution et Mediterranean the tropical Moharrek in the the 2017; occur in of Borsch, also Genera diversified & genera Ta- Limonoideae. initially few the Supplementary have a and and to hough Plumbaginoideae The thought the 1 Polygonaceae). subfamilies, are (Figure and two 2003). Limonoideae Caryophyllales Plumbaginaceae of and al., Tamaricaceae, comprised the et Nepenthaceae) (Frankeniaceae, is Hilu Droseraceae, of Plumbaginaceae clim- taxa 2002; Drosophyllaceae, clade rarely sister al., (Ancistrocladaceae, Polygonids non-carnivorous tress, et carnivorous small other Cuenoud the comprises (e.g. or which in monophyletic shrubs 1), included as herbs, ble supported is perennial well family of is The family that cosmopolitan 1993) (Kubitzki, a bers is Juss. Plumbaginaceae 2015). The noncore- al., et the Hernandez-Ledesma (Brockington Ancistrocladaceae, include 2009; Plumbaginaceae Nepenthaceae, al., to and et Polygonaceae, Drosophyllaceae, Frankeniaceae, Centrospermae) Tamaricaceae, Droseraceae, allies original families and the Dioncophyllaceae, carnivorous the with - essentially Caryophyllales corresponding (i.e., Caryophyllales Armeria Rumex Aegialitis Limoniastum Goniolimon , .maritima A. and r attlrn re mnrvs Akno ta. 97 a,2002) Das, 1967; al., et (Atkinson (mangroves) trees tolerant salt are Rheum Myriolimon lmao .auriculata P. Plumbago, , r aohts(aaa lNga,&Rmdn 99 Zouhaier 1999; Ramadan, & El-Naggar, (Salama, halophytes are .tataricum G. sfudi atmrhs(..Rzm,Gd,&Plak 1981), Pollack, & Gude, Rozema, (e.g. marshes salt in found is Plgnca) h otcmo oltr aeasakwith stalk a have colleters common most The (Polygonaceae). Psylliostachys , Aegialitis .diffusum M. 2 Ceratostigma Frdy&Tosn 96;Pwok,16) is 1963), Pawloski, 1986c; Thomson, & (Faraday steol eu fteLmnoda iha with Limonioideae the of genus only the is Put&Rvl 92 and 1982) Revel, & (Pount Frdy&Tosn 96;Frdy& Faraday 1986a; Thomson, & (Faraday ( .spicatus P. , .plumbaginoides C. .Tegenus The ). Brhr,1989), (Borchert, Limonium .ferulaceum M. has Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. ihgad,woesl oeac a e ob salse;frexample, for established; be (see of to glands presence yet salt the has utilise (e,g., where al., tolerance all established but et salt that habitats/environments, be likely (Salama whose saline leaves to is spikes glands, in it on yet and thrive with family (inflorescences) has that 2); the scapes some (Table glands in rachis, are Plumbaginaceae functional species there as the halophytic although such the of 2A), organs, Among Table genera aerial 1890). nine other Wilson, as in 1999; well described structures, as epidermal been stems, are have (Flowers, and origin glands glands evolutionary salt salt multiple cases, their Multicellular all to point In 2010). that Bromham, dissimilarities 30/Oct/2019). & structural (recreto- eHALOPH 21% Galal, and halophytes Poaceae, in anatomical remain- salt-secreting data 28%; with the with containing of but (Plumbaginaceae, families families (analysis species seven 12 just with 10% the 6%) in the Frankeniaceae, ing of and found Of 90% 15% are Tamaricaceae, approximately halophytes. 20%; they Amaranthaceae, contain contain uncommon: families that relatively five families are halophytes), 111 glands the salt-secreting of mucilage, 12 secrete that glands Unlike vascular these without 2). evolved GLANDS of have and SALT glands 1 delivery of 3 other of (Tables indicator mucilage, the glands salt secreting an favour secrete (e.g. glands not vascularized that from to is not Apart glands vasculature are seems connections, 2013). of com- species Specht, phloem carnivorous presence the & Renner of near of the ; glands However, many localisation many of 2017). since pathways a carnivory, al., synthetic so functional et the (Sharifi-Rad and in involved compounds phloem. 2009) be different the associated (Evans, through struc- to are precursors known secreted secretory of Cannabaceae, are internal supply pounds and itself regular Many photosynthesis a Clusiaceae requires and Papaveraceae, synthesis cells. the al., Plastids Euphorbiaceae, compound adjacent et and since of between Sharifi-Rad bundles, strands ducts vascular 2013; substances vascular with and to Zhu, of special & glands modified with transport Liu like been Yuan, supplied the tures, (Guo, have are regulate plasmodesmata that secretions which wall leaves producing cell 2017), with numerous glands show associated the cells are of surrounding Some glands 2018). multicellular prey. Hiscock, non-carnivorous & specific capture of Bauer, plants, glands (Thorogood, carnivorous attacks structure calyx In predatory gland The preventing ancestral so 2016). common ants, a Haridasan, like suggesting predators & , insect (Panicker for pollinators barrier in & function a insect Bharuth lying exudates Plumbago as sticky Naidoo, numerous, flying acting the (Singh, very favoured by that flowers hypothesized pollination are of on been to calyx and has aid and 1890). sepals, It an petiole 1890). (Wilson, and as the Wilson, cells on bracts, 2019; mucilage Sudhakaran, arranged laminae, sticky 2019; regularly a on Baijnath, by secrete leaves, bounded that the hairs depression, of glandular circular axils or the oval in an of cell- found species the in are at in than cells colour as larger type black but In same number, a 2B). the in (Figure of of vacuoles as appearance are epidermal trichloride well glands the few iron as mucilage a by and inside 1981) polysaccharides) acid and of Fahn, (non-structural level tannic epidermis and wall mucilage lower test, (Trachtenberg the of the histochemical protoplast at on presence mucilage either performed and A the 1973), accumulate wall from 1988). Lillie, may (Fahn, cell radiate & gland mucilages cells between and (Pizzolato the The epidermal covering space conical 1890). of envelope or the Wilson, vacuoles cuticle columnar 1916; in in 1992; the Fraine, Fahmy, prismatic, or in de viscous are & level pores 1992; colourless, cells Hassan, cell-wall without al., transparent (Batanouny, walls, secreting et a periclinal surface The (Batanouny of straight (adaxial) petioles. cells quantities with large the upper cells secrete of collecting its and base basal on 2A), some the sheath Figure In at leaf 1890; organs appearance. liquid Wilson, the other in on 1916; of similar Fraine, and relatively leaves base de of are the axils they at the where in occur genera, appear mucilage-glands the These all (1916). of Fraine de and (1890) Wilson Aegialitis , r ntmclysmlrt h uiaegad fcrioosgenera carnivorous of glands mucilage the to similar anatomically are Armeria , Ceratostigma , Limoniastrum Limoniastrum Limonium yilmndiffusum Myriolimon , Limonium 3 u in but , Wlo,19) In 1890). (Wilson, Armeria rei,Ceratostigma Armeria, ioimmultiflorum Limonium and Plumbago h aa el r oprtvl few comparatively are cells basal the al ) hr r lospecies also are There 2). Table ; lmaozeylanica Plumbago Aegialitis Limonium ncmrhniesuisby studies comprehensive in Drosera h mucilage-secreting the , Plumbago and pce,teeglands these species, evs demonstrates leaves, Limoniastrum and Drosophyllum (Sudhakaran, pce have species Nepenthes the Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. aaon ta. 92 aaa hmo,18c aaae l,19;Vsiyv&Seaoa19;Yuan, 1990; Stepanova & Vassilyev 1999; al., et Salama 1986c; Thomson, & Faraday cells 1992; 16 al., reports et Batanouny (1986c) Thomson and material. part Faraday In published central by the a an study from by and number the surmounted cells, either B-S Although one confirm accessory Wang, each 2015). cells, six 1967; collecting al., cells, al four for et collecting et - (Zouhaier huge cells (Atkinson two cell eight cells containing including apical eight structure 4B), secretory of (Figure the rings section representing Salama three cross 2014; consist in al., and 16 et 2002) In Feng (Das communication). 1986c; layer personal Thomson epidermal in layers, & whereas other the Faraday two 1986c) in of by Thomson 1988; leaves enclosed and (Fahn, (Faraday In being cells cells both four 1999). cells, of al., accessory consisting by et surrounded them are of cells each secretory central the four ‘accumulating collecting in The cells).. cells, and included transfer secretory cells, are terms or collecting the (=accumulating cells In use to cells basal and accessory refer synonymize large we and to four simplification cells) cells’ For not (=basal There ‘basal cells. or cells 4A). accessory term whether to characteristically (Figure the refer to 1988), structure use to as cells’ al cuticle-lined authors & literature et different Balsamo a the 2; that Thomson in in (Table gland11Note 1999; cells, disparity cells al., some accessory 16 however, of et and consisting is, Salama collecting family secretory, 1986c; the into Thomson, within differentiated & species arrangement Faraday most cellular 1993; of this glands Thomson, the reported with also history preserved, a (1915) well for Ruhland 2017 maritima and structures). Armeria 2016 four glandular Toma, in and these arranged Grigore of cells in (see (with description Plumbaginaceae glands the the salt within 16-celled of found complex 4) the of of Figure leaves description quadrants; of anatomical sections detailed longitudinal a and illustrated cross using (1915) having Ruhland as gland recorded per is cells species no of mm Plumbaginaceae, number of per mm group smaller 12 per sister of This Polygonaceae, glands. 31 (median the salt Plumbaginaceae of 3). of the (median members and than in and in frequency rather seen Frankeniaceae 2 Notably, higher than 4), (Tables the (Table 4) a Plumbaginaceae within Table eight with Tamaricaceae, the generally Glands associated - have within is Plumbaginaceae Tamaricaceae species 2019). the (8) the the October in salt- within genera in (eHALOPH, of species within than 80% 16 glands halophytes salt-tolerant cells, halophytes, of salt fewer of all number have having of 2017; 67% the Tamaricaceae respectively, as Toma, Although and 1%, Frankeniaceae recorded and 4). and (Table the Grigore been 2% species within just 2010; 58 small, species and al., is tolerant genera et Tamaricaceae four Flowers and species, in and Frankeniaceae Frankeniaceae - 1988; most the families 1988) of Fahn, Within Caryophyllales al, glands noncore 2017; et the the cells. Thomson Larkin, in only 40 described & 1988); been (Dassanayake to Oross, also Tamaricaceae up & have Faraday, glands of (Thomson, salt consisting Multicellular cells complex, 16 of are guyonianum composed Plumbaginaceae are the glands of in the the glands glands within genus salt salt largest to The the analogous in common structures particularly glands,). of is salt presence established with be The to 2B). yet (Table has family, tolerance glands salt salt where to species similar structures and have 2005) Perez-Garcia, & Escudero, and 2019) ttc incana Statice Armeria Limonium lmaoauriculata Plumbago . opooia tde fsl glands salt of studies Morphological 3.1 Limonium .europaea P. , wt 2cls als2ad3adFgr )adtoeof those and 3) Figure and 3 and 2 Tables cells; 32 (with Ceratostigma pce ope 6cle atgadsrcuei omnyrpre Tbe3 iue4A; Figure 3; (Table reported commonly is structure salt-gland 16-celled complex a species (synonym: a es 4seis;seTbe Aad3;teeae3 aohtcseisof species halophytic 34 are there 3B; and 3A Tables see ; species 14 least (at as hnkonas known then (also Wie,17)a ela o-aohtslike non-halophytes as well as 1972) (Waisel, n hto uhkrn(09 of (2019) Sudhakaran of that and eiltsannulata Aegialitis and oilmnincanum Goniolimon ioisrmguyonianum Limoniastrum Goniolimon .canescens A. .vulgaris A. h 6clso h atgad r ragdi orquadrants. four in arranged are glands salt the of cells 16 the atgad pert eognzdit ih ig ffive of rings eight into organized be to appear glands salt , .rotundifolia A. ) .latifolia S. , Saslai aqa alta bae 06,which 2016), Abbate, & Valletta, Pasqua, (Scassellati, 4 .Ide ntePubgnca hsognzto is organization this Plumbaginaceae the in Indeed ). evs oee,ec atgadcmrss3 cells, 32 comprises gland salt each however, leaves, .zeylanica P. hyaepeetwti u-hpdcrypt cup-shaped a within present are they (synonym: ttc gmelinii Statice 2 rei caespitosa Armeria 4wti h rneica and Frankeniaceae the within 14 = n ; hw ih el,i sntpsil to possible not is it cells, eight shows Aegialitis ioimplatyphyllum Limonium (syn. 2 r4 el)hv more. have cells) 40 or (24 2 =2,Tbe n 3). and 2 Tables 22, n= , (Gimenez-Benavides, ioimgmelinii Limonium Limoniastrum Limonium n in and ) ) Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. otlrt aiiywtotteaiiyt xrt atwieohr tls lns r hs lnsrequired able glands are these species Are some how glands. of utilise question others a while raises salt halophytes excrete all, to ability not saltmarshes. the but Kalogerakis maritime of without some, & plants on salinity in Manousaki, while growing tolerate (Kadukova, glands plants 2017), to salt Cd of al., of and secretions et presence Pb (Santos the The Zn dominate secrete and Cl to Pb and shown salt Ni, Na the been Na, Nevertheless, of have Mn, 2008). cells culture Mg, within solution K, Zn Fe, in identified Cu, of growing (2002) Cr, leaves Heumann Cd, of culture, Ca, surface solution As, the in on plants Salts In glands. 1995). al., et (Neumann of leaves Na of fluxes with 99.Wietescein rmmn pce ocnanC n g(al ) h lnsaecpbeof capable are glands the 2), al., (Table et Mg Salama and 2014; Ca al., contain et (Na Feng do ions 1986a; species of Thomson many variety & from a (Faraday the secretions secreting glands revealed low the by have the microscopy, While secreted x-ray on are 1999). analytical based that suggested particularly structures” elements techniques, (1974) secretory of analytical Sakai separate range in indeed we not Advances and those are that substrate. 1890) glands to way on Wilson, salt the similar by and and structures shown glands 2017 of selectivity chalk Toma ion are probability & illustrations all Grigore Their “in (compare that glands 1870). (Wilson, salt (Licopoli, ‘Mettenius’ call 1886), glands (Maury, now ‘Licopoli’ ‘calciferous’ called even and (1866) or Licopoli 1890) and (1856) Mettenius by drawn were CaCO of secretion 4). the (Table , century, cells nineteenth eight late totalling the cells In secretion collecting two salt and of arbitrary cells Physiology most an secreting showing 3.2 six be as of to data comprised are us the being exceptions interpret to each We the with appear gland. glands; cells the 16-celled guyonianum 20 in have Limoniastrum and included Plumbaginaceae are 12 the E cells between of 5 which members number (Figures 1988). of (Fahn, cell deposits interpretation a bundles wax in of vascular gives of difference differences the which types with 5D), reported secretory connection different (Figure summary, four direct have cell) In The no can per have cuticle pore 1969). glands whose (one The al., epidermis, pores F). et visible the and (Thomson four to damage through appearance salt solutions peculiar from mineral cells the of excrete mesophyll sections cross cells and of on view epidermal of reported microscopy surface in protecting as fluorescence insertion on observed by structure in the and cortical confirmed as elevated 5B) is spikelets, surface special which (Figure and the a gland, leaves of the scapes from top forming protrude stems, the cells may on In of glands located & with be Gharibiyan 1988). leaves: side even the Mahdavi, al, in the or by on Malekmohammadi, 2001) side in et Lefebvre, seen Akhani then that Thomson & be B; being (Bernard to leaves, 1999; can & similar the glands 5A al., is in the Figures et sunken glands leaves, 3, deeply Salama In and or 2013; cells 2 epidermis. Chase epidermal (Tables the other cells in collecting the mesophyll position basal of the a its level 5A). the in (Figure cells: at vary cell four surface secretory can are a cells complex there surround of glandular quadrant, these number The each and the In cells, that accessory differentiation. that possible two functional show and is and we cell it spatial Here however, collecting a cases, four have interpretations. and previous cells different cells, in to accessory As open four nydeggeri is 2012). and gland internal 2011, a four In Chu, by within & 2012). bounded Tan Shen, and (Xin, found. & cell cells Tan been adjacent (Ni, an also cells) by have collecting accompanied cells and of adjacent arrangements secretory, franchetii other central although (four cells 1966), 12 Luttge, & in Ziegler instance, For 2016; Wang, & Leng Goniolimon ioimpruinosum Limonium .maritima A. and h auesl ln sdsrbda ope tutr ih2 el orsceigclseach cells secreting four - cells 20 with structure complex a as described is gland salt mature the , .ovalifolium L. Limoniastrum .aureum L. + n Cl and rwn ncnaiae olcnandC n esraonso n i eadMn and Fe Ni, Zn, of amounts lesser and Cu contained soil contaminated on growing Slm ta. 99.Adneadtikctcei rsn rudtewoegroup whole the around present is cuticle thick and dense A 1999). al., et (Salama - , ecigvle foe 0 mlgadh(al ) rsaso h ufc of surface the on Crystals 3). (Table pmol/gland/h 200 over of values reaching pce ihnteFaknaeeadTmrcca aeegtcle glands eight-celled have Tamaricaceae and Frankeniaceae the within Species . .bocconei L. l rsn atgad ih1 el,arne nqarns(iue5,where 5), (Figure quadrants in arranged cells, 16 with glands salt present all and + K , Plumbago + Ca , aai aphylla Tamarix aai africana Tamarix 2+ , .lojaconi L. .narbonense L. Mg , a eosrtd(rcno,13) h rasresponsible organs The 1836). (Braconnot, demonstrated was 2+ Zn , 3 5 n in and ygad ntesraeo evsof leaves of surface the on glands by 2+, a rdmnnl ert ihrN rC depending Ca or Na either secrete predominantly can evs(iue5) hssest efundamental be to seems This 5C). (Figure leaves Cl rwn nacnaiae atmrhcontained marsh salt contaminated a on growing - HCO , .pignattii L. 3 - SO , .multiflorum L. h auegadsest include to seems gland mature the 4 .anlt,A rotundifolia A. annulata, A. 2- e loTosne l 1988) al, et Thomson also see ; , .sinense L. .nroes L. narbonense L. Armeria .multiflorum L. .smyrnensis T. .maritima A. , Statice and and L. Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. lentv xlntosfrtepeec fgad nmmeso h aiyado pce ihnthe within for species looked of have and we important family an below) the is significant and Ca of a concentrations. 3B members Ca is 2B, of in there regulation (Tables yet the glands halophytes is and of possibility not glands One presence are salt Tamaricaceae. the and utilise that Frankeniaceae for to glands appear explanations with Plumbaginaceae alternative species the to species. of in linked some not species number in secretion also tolerance Na tolerant compares salt is of salt for that concentrations As . required halophytes xylem . . are for the of sufficient glands glands data or study not not salt systematic uptake is or of a no trait) whether of presence In is gland The proportion there salt survival. Unfortunately, a tolerance. (the for salinity as excretion exclusion”. critical of salt salt levels be of . high could . levels for confer “. proportion to content that small necessary concluded in or (2015) a increase Lovelock even of & but NaCl Reef proportion mangroves, mM secretion, a 200 on as in reliant an growing excretion particularly plants be calculated For can 1981) glands species. al., by maritima, some too excretion et of is that (Rozema shoots there showed anglica in they but Spartina (1981) load excreted, days, al. are salt three et ions the Rozema for excess balancing in if conclusion. of mitigated Cl clear aspect or a and important avoided 2015). allow Na be Lovelock, to utilise might evidence & situation halophytes little Reef This many reduced. that 1985; values Na is fact (Flowers, growth of exclusion’ the % delivery Nevertheless, by Na 99 the of for to time. concentrations once accounted time xylem 77 that over be that from in content can so range transpired shoot growth exclusion’ and in in water ‘low growing change calculated of apparent are from volume The been plants estimated have the Neither if be by halophytes medium. easy of to poorly the divided for only have Estimates a in weeks) are concentrations have those to concentrations xylem that to rice 1987). (hours medium sap while system and Flowers, xylem medium root determine: process the and the liquid to in this - of Yeo a easy concentrations 2008) at areas Yeo, ion particularly Tester, through poor comparing is 2010; & ions by rather parameter Flowers, (Munns of made be leak extent be & to can a some Al-Azzawi ‘exclusion’ flow; appear to (Faiyue, plants ‘bypass’ rice, vessels all a endodermis as xylem that have developed such clear the to is reaching shown species, It been ions Salt-sensitive rate. has of growth exclusion. entry their called the with ions so restrict delivery leaves? ions to the of the able balance access reaches to the are that have limiting salt plants genera in salinity, excrete Amaranthaceae the survive to the To have of example, members so species for the and that than xylem in, it efficient the Is less euhalophytes essential to Tamaricaceae? are not are, and families Frankeniaceae clearly three there Plumbaginaceae, these are in the since in (present again of Poaceae members Poaceae, but the euhalophytic 2017), the of of feature Larkin, microhairs in The & salinities Dassanayake do. Phragmites (see high Poaceae Paspalum, salt the of of secrete tolerance glands do in the for the Pooideae) tolerance that in the salt way seen extreme the but cells for in all as essential storage ions such not the excrete genera are to not in bladders analogous do absent Such the system, are family. of they salt-storage the Those Amaranthaceae; a of the families. as members other other interpreted Bose, the in be Shabala, leaves in 1969; can succulent seen Shacher-Hill, and & glands 2014) Pallaghy Amaranthaceae complex West, the Hedrich, the Luttge, of & glands (Osmond, from salt bladders different The external so are glands? and Amaranthaceae and salt Frankeniaceae bicellular of related (Frankeniaceae, are importance high the the Poaceae remarkably why In also and So, is Amaranthaceae. salt most glands 60%). have the Tamaricaceae, salt euhalophytes in the and with 18% its 100% species amongst and of of euhalophytic Poaceae of are (86%) 80% are the proportion most for so of the species that accounting Tamaricaceae, 31% and 74 Plumbaginaceae in to just remarkable and salt compared however, which Poaceae have as mM is, of Amaranthaceae, glands and Plumbaginaceae 200 the eHALOPH, The reasons with in halophytes, least - species. listed extreme other at glands these euhalophytes more for salt tolerate 524 the evolved have are can for to that There case that reported structure the plants. plants examine of a are we species simply Here these salt-tolerant they present? - are when euhalophytes salts or the secrete species to some adapted in become tolerance salt for .4 ti la httetommeso h lmaiaee( Plumbaginaceae the of members two the that clear is It 0.04. ob ,for 1, be to and + n Cl and Puccinellia ioimvulgare Limonium - xed h aaiyfrtepatt oprmnaietoein,then ions, those compartmentalise to plant the for capacity the exceeds htd o aesl lns o h r atgad uhacommon a such glands salt are why So, glands. salt have not do that + a ehg se o xml,Yo&Foes 96.However, 1986). Flowers, & Yeo example, for (see, high be can h ai a .;for 0.3; was ratio the , 6 Salicornia + n rCl or and , Suaeda lu maritima Glaux Limonium or - htwudalwu oevaluate to us allow would that Tecticornia and . n for and 0.1 , Armeria n,weepresent, where and, eenot were ) Armeria Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. l,21) h ln hrce sfudi pce agn rmprnilhrswt lgtyflsylae as leaves fleshy slightly with et herbs in (Moharrek perennial in rocks from in exposed ranging present as on species trees not or in found soils are is stony glands character and in gland gravelly although Tomovic, The on & 1972), habitats 2019). Jakovljevic dry al., Waisel, Niketic, in Stevanovic, 1986c; (Buzurovic, regions mountainous stems Thomson, and colonized leaves & its Faraday on glands 2013; salt in has (as regions on spray hilly grow salt that genera airborne in in of present and one deposition also 2014) are to al., Glands exposed mangrove et 2011). soils Caperta the Liu, incipient and (including in Zhao areas Feng, coasts Song, rocky tidal Zhao, in 1916; Fraine, grow de can 1995; 2015). that Savoure, species & in (e.g., Flowers occur Bouchereau, glands arid Abdelly, glands) Aegilatis salt multicellular to Slama, adapted Plumbaginaceae, (e.g. compounds; anatomy species the involving osmoprotective and adaptations In genera – (e.g. habitats with physiology saline tolerance, of and drought these range and wide of a salt species and for environments allowed well-known this is time, Plumbaginaceae The of glands ( course seawater salt the as of Over well Evolution as 2017). 3.4 soils in Luan, saline the evolved & drier perhaps have (Tang colonise and could to Mg concentrations glands Ca families and salt shoot Ca multicellular regulate between that to balance Tamaricaceae hypothesise of and We Frankeniaceae shoots Plumbaginaceae, regulated. the the for be of value could Ca The concentrations shoot both Ca literature. the of shoot the glands recorded of in the 2003) 4.41% data from to al., from 0.11 et is (2003) (Broadley seen plants values al. they in seen of et concentrations range Broadley Ca lower ramosissima by of shoot Tamarix the calculated however, range although in The aware, as Tamaricaceae, be weight are 2003). or to dry Frankeniaceae, al., we appear et Plumbaginaceae, Tamaricaceae, as (Broadley the and far plants Plumbaginaceae of across As the relations Amaranthaceae in Ca 2003). the concentrations Broadley, the the Ca in of & is particularly which known (White seen amongst is Caryophyllales 2017), 2005), little the Luan, Nakata, & within & (Tang (Franceschi Polygonaceae concentrations oxalate Ca and there calcium their Since plant regulate precipitate molecule. for to to signalling means essential ability important of Ca is variety an for a as Ca force role use plants its driving 2012). reflecting inward effects 2017), Karimi, strong Ca its Luan, free a & through & cytoplasmic is Tang Hadi also of 2003; concentration but 1980; al., the signalling, yet et Munns, in and Broadley role & 2003) Broadley, its (Greenway & in membranes (White only growth and not salinity, walls to cell plants on of response the of determinant tngt(uie,Rbn,&Je,18;Rne pct 2013). Specht, vapour water & of Renner absorption glue, 1989; the trapping contain as Joel, generating well may & lures, as and Robins, as medium such pitted, (Juniper, (enzyme) functions digestive or night different a at stalked, of and sessile, variety mechanism a drowning be evolved a can have providing subfamily they glands the phloem; multicellular than and the xylem structures 3), of (Figure variety families greater carnivorous families, a pos- non-carnivorous with likely related and Plumbaginaceae the subfamily Limonioideae glands family in the salt present the Plumbaginaceae, members with also the of phyloge- salt-tolerant genera are Within ancestor Our in glands Tamaricaceae. common absent such and recent vasculature; understood. are Frankeniaceae, without most well glands glands the salt not salt Second, First, are sessed syndrome’ Polygonaceae. findings. glands the main ‘salt of several the revealed reconstruction to netic leading pathways evolutionary & The (Baumbach regions geographical different in 2007). independently Hellwig, and repeatedly populations non-metallicolous Limonium .maritima A. Goniolimon Limonium a,Msr,&Mhny 06 n nhge re at fsl ase ujce ohg salinity high to subjected marshes salt of parts drier higher in and 2006) Mohanty, & Mishra, Das, ; Aegialitis pert aedvrie oeadmr eetyta the than recently more and more diversified have to appear ocsinfrigdafsrb sin as shrubs dwarf cushion-forming to , rvd togeiec htmtlioosppltoshv endrvdfo h ancestral the from derived been have populations metallicolous that evidence strong provide and pce httrvsi tpelk aias(eohlu atrsadrcygons in grounds) rocky and pastures (xerophilous habitats steppe-like in thrives that species Tbe ) ugsigapeimrhcoii fti aohtctat eei studies Genetic trait. halophytic this of origin plesiomorphic a suggesting 3), - 1 (Tables Limoniastrum si h ideo hsrne t19%o h r egt nahdooi experiment hydroponic a In weight. dry the of 1.97% at range, this of middle the in is .ramosissima T. Armeria lae aznrs 07 ot ta. 04 asn&Ingrouille, & Dawson 2014; al., et Costa 2017; Manzanares, & Alvarez ; pce fcatlsn ue Asn izGrea,21) tleast At 2017). Diez-Garretas, & (Arseni dunes sand coastal of species 2+ and nocls(eici,Saaa syno,Ci otsn 2018), Pottosin, & Cuin Isayenkov, Shabala, (Demidchik, cells into .maritima A. Armeria 7 rei maritima Armeria Aegialitis Tbe n ) hsi en ywihshoot which by means a is this 4), and 2 (Tables osa husa in as shrubs coastal , al 2). Table ; ob .9.SneC ssecreted is Ca Since 0.59%. be to Plumbaginoideae Limonioideae Limonium, 2+ Limoniastrum slw(sub-micromolar, low is Plumbaginoideae Acantholimon nterelated the In . rcyspecies’; ’rocky rsnsmore presents n small and the : that Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. 03 aaae l,19;Wiln&Wne,19;Zage l,2018). al., et Zhang 1991; Winter, & Weiglin (Kohl, 1999; populations al., as marsh et such salt Salama as 2013; than halophytes such (NaCl) and non-halophytes water both 4) saline Tamaricaceae, Table with related the irrigation (e.g. In to pastures resistant 1997). mountain less and if meadows even sub-mountain (e.g. salt-tolerant and in soils 3 and sandy 2, 1915) Tables Ruhland, non-saline canescens (see In 2007; in ions Hellwig, clarified. innovation found of & be ecophysiological range metalophytes Baumbach to an a yet and as is secrete halophytes evolved environments to saline glands in adapted in salt and carbonate those Whether above) including and (discussed 2017). 4) mudstones concentrations al., and Ca et conglomerates regulate sediments (Caperta sandstones, on to limestone thrive like (Ireland, typically in rocks Europe species as sedimentary in these rocks coasts from that Atlantic revealed derived the over Morocco, deposits all in and complex, and each Peninsula) from Iberian species France, the Britain, of range distribution in entire the as arid cliffs, in rocky and as L and rocks, harsh marshes exposed of potentials of and coastal diversification water colonization soils ovalifolium steppes, low stony the allowed saline of tolerated facilitated that from xerophytes plants have halophytes and adaptations might and example, anatomical This for ages and conditions; previous physiological 2006). in Callaway, possessing than & species drier Verdu, took were Rumebe, separation conditions (Valiente-Banuet, this climatic that the is split when the it upon how place but took Plumbaginaceae, Limonium glands salt group with sister lineages unclear. its of is and establishment place the Polygonaceae that the presume between might we of halophyte up, sum the To with . together clade a by form glands with Ceratolimon also subfamily. species the within non-halophytic genus Psylliostachys and oldest glands the be with to species seems major glands, two salt present Plumbaginaceae, members the Within Plumbagella 2). the Table of Supplementary subfamilies 3; monophyletic (Figure Plumbaginoideae two the outgroups tRNA-Leu represent as chloroplast clades used ( using Polygonaceae here Plumbaginaceae the the ), of members of salt-tolerant analysis sister inference from Bayesian split a the revision, ( placed 2019). analyses current al., Molecular the et In Ma. (Moharrek 57 Pliocene – the and 27 Miocene at Late Plumbaginaceae Tamaricaceae the of for age ages the non-halophyte maximum to and minimum compared sister dated Schuster, Ma its (2010) ago, between 58 and Soltis years Polygonaceae – & (million the Soltis 60 Ma between Bell, 90.7–125.0 at split about 2013). the Kron, occurring that & ancient, estimated Setaro, relatively been is has Plumbaginaceae it the dating, group molecular on Based Plumbaginaceae no as ble. to the are 1984) referred Ghobary, from there Weber-El are microfossils since 1976; and of Nowicke, are estimate distinguish those Tamar- there to to grains, related difficult Although difficult pollen and are is of families. Plumbaginaceae form families these of the these of members in of by macrofossils date shared reliable divergence are recorded the glands Frankeniaceae, salt and icaceae that fact the Notwithstanding trnL . Acantholimon multiflorum eeand gene ) Ceratostigma Plumbago cselt ta. 06.Iln ouain of populations Inland 2016). al., et Scassellati ; pce emdt iesf tafs aedrn h esna Leoe l,20;ti study), this 2005; al., et (Lledo Messinian the during pace fast a at diversify to seemed species ( .lneltm .ndgei .ovalifolium L. nydeggeri, L. lanceolatum, L. n ls to close and , ntesubfamily the In . cnhlmnsnulato sensu Acantholimon nte ld ihhlpye ihsl lnsi htgrouping that is glands salt with halophytes with clade Another . opee n in and complexes ) , trnL-trnF , Dictyolimon Plumbago and ld,wihaanhshlpye;tgte hyfr ld ihtemonospecific the with clade a form they together halophytes; has again which clade, Limonium Myriolimun negncsae,sosta eeawti h lmaiaeeaeseparated are Plumbaginaceae the within genera that shows spacer, intergenic eu ihhlpye ihsl lns scoeto close is glands, salt with halophytes with genus a , , Bukiniczia Limonioideae Reaumurina .algarvense L. ewe 8–1 adrn h icn Leoe l,20) hra the whereas 2005), al., et (Lledo Miocene the during Ma 16 – 18 between Acantholimon n to and iegdfo t itrgnsaot75M,bgnigbtenthe between beginning Ma, 7.5 about genus sister its from diverged . ec,etmtn iegnedt sn oslple sntfeasi- not is pollen fossil using date divergence a estimating Hence, , Chaetolimon Ikonnikovia and h genus the , sn curnercrso ufc ihlg encompassing lithology surface on records occurrence using Limonium Tamarix Plumbaginoideae , 8 Armeria cnhlmnsnulato sensu Acantholimon Polygonum ld ihnnhlpye ihP= sformed is PP=1 with non-halophytes with clade A . .maritima A. , and ) Vassilczenkoa or pce oss atgad Abuzs tal., et (Abbruzzese glands salt possess species Limonium, Armeria , .binervosum L. Goniolimon Armeria, yiai germanica Myricaria P=)ado aaiaee( Tamaricaceae of and (PP=1) ) and rmsnyadhaymtlsisare soils metal heavy and sandy from .vulgaris, A. ye(ae,14,15;Svra& Skvarla 1953; 1948, (Baker, type ihmn aohtsadwhose and halophytes many with , Limonioideae ertr lnsaepeetboth present are glands secretory Popoviolimon , Limonium ( Armeria td nthe in study A . .bnrou,L dodartii, L. binervosum, L. Dyerophytum Limoniastrum synonym Dre ta. 2017; al., et (Dorken P=) niethe Inside (PP=1). ihahalophytic a with , and and Cephalorhizum .maritima A. Psylliostachys n sister and , Limonium Limonium itrto sister , Tamarix A. ; Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. h usinw aentbe bet nwri hte o-ertn lnseovdt ert aland NaCl Ca secrete to regulate (e.g. evolved to soils glands evolve saline ion-secreting glands on whether ion-secreting the is Na if answer in of or to secretion tolerance salt able tolerance salt with been for salt species not essential generated halophytic have are where of we glands species proportion question ion-secreting salt-tolerant high The that some The view as salt investigated. well the for be as supports to critical salt Plumbaginaceae. family yet is to the has uptake tolerance functional within Furthermore, glands untested of to glands eHALOPH). salt similar with proportion structurally of in but glands small presence included have species, a the that other species demonstrable family of in the 54 have excretion within glands the that species even salt are of species that there of (46 Nonetheless, suggests across glands majority available tolerance. glands the salt data mucilage Plumbaginaceae, than functional limited to common the have rise the gave less For glands salt, former are the the to glands plants. multicellular whether of salt flowering tolerance unclear are However, structure is of independently. glands it the families arose glands, these Although they salt the All of if that uncertain. or from those latter is ions. distinct between the ancestry is inorganic differentiated materials their secrete be organic but secrete can that that plants, plants those of dicotyledonous and parts in aerial compounds structures the organic on secrete evolved have that that structures between secretory connections The known. direct yet However, not are 2018). Plumbaginaceae Conclusions al., the 4 et in Caryophyllales, of (Yang innovations non-core propensity biological inferred the the and were within leading show duplication events (Rivadavia, and branch transcriptomes, genome paleopolyploidy Caryophyllales, Droseraceae on the six of carnivorous based along least to history sampling or at evolutionary leading phylogenomic 2013), branch the A Ogburn, Edwards, the throughout 2003). & & and polyploidy Hasebe, (Edwards 2013) Ogburn & succulence al., 2008; Kato of et Edwards, Kondo, evolution (Schuster & the Polygonaceae paleopoly- Ogburn with the the as Eggli, associated to such Portulacineae , 2014b), Peer of Nyffeler de base Van 2012; 2016; & the Soltis, Maere Vanneste, at & Soltis 2014a; event Peer, 2015; ploidy de al., Van et & (Edger Maere paleopolyploidy Baela, Vanneste, with correlated been have innovations Evolutionary otcmol,sl lnsi h lmaiaeeaefre rm1 el.Weegnr aegad with glands have genera know Where not Na cells. like secreting do 16 of we cells, from capable data formed 16 are glands of than Plumbaginaceae have lack the more they could a in if glands glands to or salt that owing commonly, salt However, view Most of the plant. tolerant to the are weight in species some concentrations gives these Ca soils if regulate non-saline to on evolved presence have Their glands). chalk (so-called sogdianum hswr a uddb otgeentoa ud hog udcopr ini Tecnologia a e Ciencia a para attributed Fundacao PTDC/AGRPRO/4285/2014 through project funds and national UI/AGR/04129/2013 Portuguese LEAF by through funded (www.fct.pt/) discussions. was insightful diagrams, work for schematic (LEAF/ISA) This drawing Abreu for Manuela (LEAF/ISA) Maria Ferreira and Cardoso Viegas Teresa Wanda acknowledge gratefully authors metals. heavy The in high glands glands soils such Salt-secreting of of colonisation presence non-saline tolerance. ACKNOWLEDGEMENTS the The in salt enabled thriving trait. have to species halophytic to in this aid maintained appears with of an albeit also family independence habitats is evolutionary a saline the structure of Polygonaceae, reflecting Plumbagi- colonization related these environments, successful the the of the in allow of evolution to glands ancestor the seem Plumbaginaceae, salt the common suggest of of halophytes, a absence members absence halophytic the few of and The in Frankeniaceae and glands possibility Caryophyllales, Frankeniaceae cells. salt closely non-core the and of eight like is Tamaricaceae prevalence evaluate halophytes just The tolerance of cannot of glands). salt families we (8-celled composed related Tamaricaceae where are that and families structures glands) means Tamaricaceae, (16-celled these record naceae and glands, fossil Frankeniaceae salt a related of of presence the the In with associated 3). (Fig. groups , .nudum L. + n Cl and rei caespitosa Armeria Aegialitis gpu soils) (gypsum - n attlrne ihntePubgnca hr r pce htd o grow not do that species are there Plumbaginaceae the Within tolerance. salt and and . Limoniastrum itdi al atB,btwt lnsta oklk atglands salt like look that glands with but B), part 3 Table in listed and , .canescens A. 9 hyeovdidpnetya hyfr non-sister form they as independently evolved they , , lmaozyaia .erpe,Limonium europaea, P. zeylanica, Plumbago 2+ ocnrtosadti enabled this and concentrations + n Cl and - . Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. ae,HG 15) iopimadmnmrhs ntePubgnca 2 olnadsimt nthe in stigmata and Pollen family. .2. the Plumbaginaceae of the survey in monomorphism A and genus Dimorphism .1. (1953). Plumbaginaceae H.G. the Baker, in monomorphism 12 and Botany, Salt of Dimorphism Annals (1967). (1948). K.R. West, H.G. Baker, & H.G.W., Saddler, 20 Science, M.G., Biological Pitman, of mangroves A.B., the 0.1007/978-3-319-54867-8 vol in Hope, https://doi:org/ Peninsula, regulation G.P., Publishing, Iberian International Findlay, the Springer of M.R., vegetation 397-432. Atkinson, The pp. In: Loid). Vegetation. Tucson Coastal J Arizona, (ed (2017). of 2 B. University Diez-Garretas, plant world. & A., the succulent Arseni, of major plants Salt-tolerant world’s M.J., (1989). the Moore, J.A. of Aronson, E., radiations Spriggs, recent R.M., Ogburn, and U., https://doi:org/10.1073/pnas.1100628108 Contemporaneous Eggli, A., (2011). Lendel, lineages. E.J. R., Edwards, Nyffeler, & P-A., Christin, M., Arakaki, dicotyledons. marshes salt Mediterranean costal from plants of 44 anatomy Leaf (1989). I.M. Vicente, relationship. Andres, communities mediter- plant & and four landscapes M., Mediterranean on 38 (2017). Boscaiu, Lazaroa, study J.A. 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Flora, 977. E. in Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. binervosum bicolor bicolor bellidifolium axillare axillare aureum eHALOPH in - tolerant salt are that species A parentheses) in (synonym Species species 3B. non-halophytic and and 2B eHALOPH Table in in listed ( listed species m ( includes 3 inflorescences $ to paniculate http://www.plantsoftheworldonline.org/. up bearing stems stems flowering woody and linear leaves, with ( ( basal shrubs inflorescences, shrubs sub- dwarf capitate or cushion-forming or shrubs and spike-like herbs mostly perennial bearing ( eHALOPH subshrub leaves database acuminate the caespitose rigid following assignments branched (“syn- type densely morphologies plant and Overall to pulvinate (2019) - Ac relationships. al. Acronyms: phylogenetic et (https://www.sussex.ac.uk/affiliates/halophytes/index.php?content=plantStats). Moharrek or on monophyly based and are dromes”) type plant form/syndrome, S. life Abderazzak, & of 1 A., glands TABLE Chedly, salt leaf B.A., of Aouatef, analysis C., X-ray and Wided, microscopy D., guyonianum electron Wahbi, transmission and T., Scanning Najla, (2015). A., Abdallah, B., Zouhaier, 10.2298/ZMSPN1325043Z. https://doi.org/ 43-54. Lukovi´c, J. Limonium & Karanovi´c, D.S., Anaˇckov, G.T., von Zori´c, L.N., salzdrusen Die (1966). https://doi:org/10.1007/bf00963728 U. 1-17. Luttge, & H., Ziegler, Limonium ilr(lmaiae,Pubgnca)taxa. Plumbaginaceae) (Plumbaginales, Miller eeaasge otePubgnca rvdn nomto ntrso ubro species, of number of terms in information providing Plumbaginaceae the to assigned Genera . os.udrNC salinity. NaCl under Boiss. o eodd1 eFraine de al. et Akhani 10 recorded Not cells secretory and cells accessory cells, cup inner cells, cup outer of each 4 cells subsidiary 4 + cells gland 8 Sunken cells basal 4 + rings 4 cells collecting 4 + 12 eHALOPH in - tolerant salt are that species A structure Gland Limonium Dyerophytum Acantholimon ioimdendroides Limonium ;L-prnilhrswt hc ottc,rslt,adsihl fleshy slightly and rosulate, rootstock, thick with herbs perennial - L ); ;A huso ml re ( trees small or shrubs - Ae ); 21 a 110 Na: 1916 Fraine de 12-16 12 9 eHALOPH in - tolerant salt are that species A mm / glands of Number irn 78, Micron, Limonium .s.l. ). 21 Armeria -.https://doi:org/10.1016/j.micron.2015.06.001 1-9. ioimvulgare Limonium 2 es lato. sensu aiaSpk ora o aua cecs 125, Sciences, Natural for Journal Srpska Matica 6N,K g Ca, al. Mg. et K, Salama Na, 2012 al. et Ni 16 16 12 eHALOPH in - tolerant salt are that species A cells of Number .(03.La tutrlaattoso two of adaptations structural Leaf (2013). Z. ˇ Limonium .C wr hus( shrubs dwarf – Ce ). Aegialitis PW lnsO h ol Online World the Of Plants *POWO Limonium .DeFeinstruktur. Die I. . agoe) r–herbaceous – Ar mangroves); ; pmol/gland/h 48-226 Cl: pmol/gland/h 40-225 SO Cl, Reference efflux and secreted elements Main pmol/gland/h ;L roecn habit, arborescent - Lw ); 4 Ceratolimon NO 3 Limoniastrum Na: lna 74, Planta, ege l 2014 al. et Feng 1999 ag2018 Wang & Wang Dong, Yuan, Leng, oa2010; Toma & Grigore 1916; 2013 ;D - Dy ); Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. multiflorum lobatum lanceolatum iranicum graecum gmelinii gmelinii gmelinii gmelinii gmelinii gmelinii gmelinii girardianum franchetii delicatulum caspium californicum brasiliense brasiliense parentheses) in (synonym Species tsrae1 hsstudy this al. et Akhani al. et Akhani 1972 Waisel 1915 16 Ruhland al. et Akhani & Faraday surface At surface At 16 10 sunken Deeply recorded Not 16 & Faraday quadrant al. et Gancedo 7 surface At rings 4 detail no but Present cells collecting 16 cells ‘goblet’ external and internal cells, secreting of each 4 cells collecting 4 rings+ 3 in cells 12 detail no but Present rings 4 cells 8 to 12+5 cells accessory remaining and cells collecting 4 cells, secretory 4 structure Gland 1Zori´c, al. et Daraban al. et Zhou Na 2012 11 al. et Xin 6-8 et Batanouny 10-12 12 20? 16 8 20-29 mm / glands of Number 22 2 6Bastos, 16 cells of Number 6ACaperta A 16 fflxReference efflux and secreted elements Main pmol/gland/h : 290 ogn 1993 Gorgen, & Perazzolo 2018 al. et Leng 2013 2013 Lukovi´c 2013 Karanovi´c, & Anaˇckov, 2013 2007 2013 1915 Ruhland, 1986c; Thomson 2017 al. et Hassan Al 1985 Blazquez 1992; al. 2013 al. et Akhani 1986c Thomson 2018 es Comm. Pers. Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. stocksii sinuatum sinense santapolense reniforme pruinosum platyphyllum perezii perezii pectinatum ovalifolium otolepis otolepis = oleifolium nydeggeri nudum narbonense parentheses) in (synonym Species virgatum (syn. ) uknAhn tal. et Akhani & Faraday al. et Akhani 16 and Faraday Sunken 10 quadrants 4 cells cells collecting 4 plus 16 12-16 detail no but Present al. et Akhani surface At cells collecting 4 16 and cells secretory 16 study this al. sections** et Akhani Cross al. et Hassan Al quadrants 4 16 12 detail no but 16 Present suface At detail no but Present surface At surface At surface At structure Gland 61 aaae al. et Salama 16 16 al. et Zhou 9 mm / glands of Number 23 2 0Xne l 2011 al. et Xin 20 cells of Number 6N:0.2 Na: 16 fflxReference efflux and secreted elements Main pmol/gland/h 20-410 Cl: Ca, K, Na, nmols/gfwt/s 2013 1972 Waisel 1999; al. et Salama 1986c; Thomson 2017 al. et Hassan Al 2013 1972 Waisel 1999; 1990 Stepanova and Vassilyev 1986c Thomson 1970 Smet Denaeyer-De 2013 2007 2002 Colombo 1989; Andres 2013 study this 2017 1986b Thomson & Faraday 2001 Alarcon & Blanco Sanchez- Torrecillas, Olmos, Morales 1980; Luttge and Jung Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. Tepeec fgad srcre ihai oun3 niae h rsneo ucinlsl glands salt functional of presence the indicates ? a 3; column in a with the recorded with is together glands makeup, of cellular presence and genus*The the frequency of database their Representatives the leaf, 3. in efflux. Table the of inclusion in by rates on indicated and glands as secreted the elements tolerance main of salt their position eHALOPH), the (from eHALOPH, glands salt have to reported 2. TABLE sogdianum serpentinicum ponzoi * pignanttii perfoliatum palmyrense lopadusanum lojaconi anfractum (syn. dictyophorum intermedium carnosum bocconei* albidum aegusae eHALOPH in not are that species Limonium B vulgare vulgare suffruticosum parentheses) in (synonym Species L. ) eeaadseiswti h lmaiaee(rmPat fteWrdOln,POWO) Online, World the Of Plants (from Plumbaginaceae the within species and Genera tsraeAhn tal. et Akhani al. et Akhani & Weiglin al. et Akhani surface At detail no but Present surface At sunken Deeply al. et Akhani Sunken eHALOPH in not are that species Limonium B cells basal 4 with circles 4 in cells 16 sunken Deeply structure Gland 0Rzm tal. et Rozema eHALOPH in not are that species Limonium B 30 mm / glands of Number oice al. Zori´c et 7 24 2 0Frdy& Faraday eHALOPH in not are that species Limonium B 20 cells of Number 6Clmo2002 Colombo 2002 Colombo & Colombo 2002 Colombo & Colombo 16 12 2002 Colombo 2002; Colombo 2002 Colombo 16 12 16 12 16 16 Limonium fflxReference efflux and secreted elements Main r rsne separately, presented are ta.1999; al. et Salama 1986c; Thomson 2013 2016 Perez Pino & Silva-Pando Javier Perez, Pino 2013 1991 Winter 2013 1981 2013 rpn 1992 Trapani 1992 Trapani 1992 Trapani & Colombo 2013 Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. Bunge Ceratostigma maritima maritima maritima Willd (DC.) Armeria rotundifolia rotundifolia rotundifolia annulata annulata annulata R.Br Aegialitis eHALOPH in Species A species and species. Genus individual than rather genera, to relate text bold in Entries known. not or uncertain is . . 2 1 glands* salt with Species 1 quadrants 4 ring per cells 8 cells 5 rings 8 cells 8 rings 3 mm No./ structure Gland a ,Ca K, Na, 2002 Das 6 ? 11-62 Cl 24 9 25 2 6NC Ruhland NaCl Bao Wang 16 & Faraday 24 40 cells of Number pmol/gland/h 0.7 Na: pmol/cm Cl / Na efflux and secreted elements Main a Mg Ca, K, Na, HCO Ca K, Cl, Na, 3 Cl, , 2 /s .9Akno et Atkinson 0.79 .6Baumeister 0.96 & Faraday 1.1 l 1967 al. Vulgaris A. (synonym 1915 1981 al. et Rozema communication personal Shan, 1986c Thomson notes & Reference 1981 Ziffus & 2013 Wang & Leng Chen, Yuan, 1989; Srivalli, & Seshavatharam 1986a Thomson ) Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. 3 Table See Mill Limonium monopetalum monopetalum monopetalum monopetalum guyonianum Fabr. ex Heist. Limoniastrum tataricum Boiss. Goniolimon plumbaginoides plumbaginoides species and Genus . 40 2 1 glands* salt with Species 24cls2 6Batanouny 16 22 cells 12+4 quadrants 4 quadrants 4 mm No./ structure Gland 2CaCO 12 & Faraday 16 31 26 2 6N,C .9Slm et Salama 0.69 Cl Na, 16 Ca & Mg 32 Balsamo Ca Mg, 16 cells of Number NaCl Cl & Na or Cl / Na efflux and secreted elements Main fwt μ 0.6-2.1 K Borchert Ca mol/g 3 , 1977 Abo & Batanouny 1999 al. soil in NaCl no when Ca & Mg 2015; al. et Zouhaier 1986c Thomson & Faraday 1986a; Thomson & Faraday 1996; Thomson & notes & Reference 1977 Abo & 1997 Ramadan 1972 Waisel icum) as (mispelled 1986c Thomson 1989 tater- ; Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. L ex Tourn. Plumbago canescens halleri ssp. maritima halleri s maritima Willd (DC.) Armeria lycopodioides glumaceum androsaceum Boiss Acantholimon eHALOPH in NOT Species B spicatus 1 Psylliostachys auriculata auriculata L ex Tourn. Plumbago ferulaceum diffusum Myriolimon species and Genus sp . . . . eHALOPH in NOT Species B ? 1 ? ? 2 glands* salt with Species quadrants 4 cells subsidiary 4 + cells 8 cells subsidiary 4 + cells 8 cells subsidiary 4 + cells 8 mm No./ structure Gland -61 Scassellati 16 2-16 Waisel 25 27 2 2Bokhari Bokhari Bokhari 12 12 12 & Faraday Ca Mg, 16? cells of Number n fflxN Cl / Na efflux and secreted elements Main ,Z Heumann Zn S, Zn Cu Ca K 1971 1971 1971 1986c Thomson & Faraday 1986a; Thomson notes & Reference ta.2016 al. et 2002 1995 al. et Neumann 1972 Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. californicum brasiliense brasiliense binervosum bicolor bicolor bellidifolium axillare axillare aureum eHALOPH in - tolerant salt are that species A parentheses) in (synonym Species tolerance. salt proven without (B) and eHALOPH) 3. TABLE zeylanica species and Genus Limonium lnua trbtsfrseisof species for attributes Glandular glands* salt with Species ig 6Frdy& Faraday al. et Gancedo Fraine de 16 al. et Akhani rings 4 cells 8 to 12+5 cells accessory remaining and cells collecting 4 cells, secretory 4 10 recorded Not cells secretory and cells accessory cells, cup inner cells, cup outer of each 4 cells subsidiary 4 + cells gland 8 Sunken cells basal 4 + rings 4 cells collecting 4 + 12 eHALOPH in - tolerant salt are that species A structure Gland Limonium tutr o/mm No./ structure Gland 21 a 110 Na: 1916 Fraine de 12-16 12 9 eHALOPH in - tolerant salt are that species A mm / glands of Number Limonium Limonium 28 2 2 A,rpre ob attlrn aeicue in included (are tolerant salt be to reported (A), 6Bastos, 16 Ca, al. Mg. et K, Salama Na, 2012 al. et Ni 16 16 12 eHALOPH in - tolerant salt are that species A cells of Number Sudhakaran 8 cells of Number Limonium n fflxN Cl / Na efflux and secreted elements Main pmol/gland/h 48-226 Cl: pmol/gland/h 40-225 SO Cl, Reference efflux and secreted elements Main pmol/gland/h 4 NO 3 Na: ege l 2014 al. et Feng ogn 1993 Gorgen, & Perazzolo 1999 ag2018 Wang & Wang Dong, Yuan, Leng, 1986c Thomson 2018 2010; Toma & Grigore 1916; 2013 2019 notes & Reference Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. otolepis = oleifolium nydeggeri nudum narbonense multiflorum lobatum lanceolatum iranicum graecum gmelinii gmelinii gmelinii gmelinii gmelinii gmelinii gmelinii girardianum franchetii delicatulum caspium parentheses) in (synonym Species virgatum (syn. ) tsrae1 hsstudy this al. et Akhani al. et Hassan Al study this al. et Akhani al. et Akhani 1972 Waisel 16 16 1915 16 Ruhland detail no but al. Present et Akhani surface At surface At & Faraday surface At surface At surface At 16 10 sunken Deeply recorded Not 16 quadrant 7 surface At rings 4 detail no but Present cells collecting cells ‘goblet’ external and internal cells, secreting of each 4 cells collecting 4 rings+ 3 in cells 12 detail no but Present structure Gland hue al. et Zhou Zori´c, al. 9 et Daraban al. et Zhou Na 2012 11 al. et Xin 6-8 et Batanouny 10-12 12 20? 16 8 20-29 mm / glands of Number 29 2 cells of Number 6ACaperta A 16 fflxReference efflux and secreted elements Main pmol/gland/h : 290 2018 al. et Leng 2007 2002 Colombo 1989; Andres 2013 study this 2017 2013 2013 Lukovi´c 2013 Karanovi´c, & Anaˇckov, 2013 2007 2013 1915 Ruhland, 1986c; Thomson 2017 al. et Hassan Al 1985 Blazquez 1992; al. 2013 al. et Akhani es Comm. Pers. Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. vulgare vulgare suffruticosum stocksii sinuatum sinense santapolense reniforme pruinosum platyphyllum perezii perezii pectinatum ovalifolium otolepis parentheses) in (synonym Species uknAhn tal. et al. Akhani et Akhani & Faraday al. et Akhani cells basal 4 with circles 4 16 in cells 16 sunken Deeply and Faraday Sunken 10 quadrants 4 cells cells collecting 4 plus 16 12-16 detail no but Present al. et Akhani surface At cells collecting 4 16 and cells secretory 16 sections** Cross quadrants 4 12 detail no but Present suface At structure Gland 0Rzm tal. et Rozema 30 al. et Salama 16 16 mm / glands of Number 30 2 0Frdy& Faraday 2011 al. et Xin 20 20 cells of Number 6N:0.2 Na: 16 fflxReference efflux and secreted elements Main pmol/gland/h 20-410 Cl: Ca, K, Na, nmols/gfwt/s ta.1999; al. et Salama 1986c; Thomson 1981 2013 2013 1972 Waisel 1999; al. et Salama 1986c; Thomson 2017 al. et Hassan Al 2013 1972 Waisel 1999; 1990 Stepanova and Vassilyev 1986c Thomson 1970 Smet Denaeyer-De 2013 1986b Thomson & Faraday 2001 Alarcon & Blanco Sanchez- Torrecillas, Olmos, Morales 1980; Luttge and Jung Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. niae yicuini h aaaeeAOH(lomre iha) h oiino h lnso the on Data glands as efflux. the tolerance of species. of rates salt than position and rather their secreted the genera elements ), ), main to a a the relate with with text with together marked bold marked makeup, (also in cellular eHALOPH; eHALOPH and (from database frequency their the glands leaf, in salt inclusion have by to indicated reported POWO) Online, 4 TABLE sogdianum serpentinicum ponzoi * pignanttii perfoliatum palmyrense lopadusanum lojaconi anfractum (syn. dictyophorum intermedium carnosum bocconei* albidum aegusae eHALOPH in not are that species Limonium B parentheses) in (synonym Species L. ) eeaadseiswti h aaiaeeadFaknaee(rmPat fteWorld the of Plants (from Frankeniaceae and Tamaricaceae the within species and Genera . tsraeAhn tal. et Akhani al. et Akhani & Weiglin al. et Akhani surface At detail no but Present surface At sunken Deeply Sunken eHALOPH in not are that species Limonium B structure Gland eHALOPH in not are that species Limonium B mm / glands of Number oice al. Zori´c et 7 31 2 eHALOPH in not are that species Limonium B cells of Number 6Clmo2002 Colombo 2002 Colombo & Colombo 2002 Colombo & Colombo 16 12 2002 Colombo 2002; Colombo 2002 Colombo 16 12 16 12 16 16 fflxReference efflux and secreted elements Main 2013 2016 Perez Pino & Silva-Pando Javier Perez, Pino 2013 1991 Winter 2013 rpn 1992 Trapani 1992 Trapani 1992 Trapani & Colombo 2013 Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. patagonica palmeri laevis juniperoides luta ( hirsuta ericifolia 11 bunda ( 11 corymbosa 10 10 capitata 78 Frankenia Frankeniaceae78 theses) paren- in onym (syn- species and Genus revo- flori- ) ) POWO species of Number eHALOPH In Species glands with Species 32 tutr Number/mm structure Gland 6+2 Sunken uknPerez, Sunken cells of No Grigore 8 2 fflxN Cl / Na efflux and secreted elements Main notes & Reference 2010 Toma & Grigore 2010 Toma & Grigore 2010 Toma & ore Grig- 1989; Andres 1991 Winter & Weiglin 1999; al. et Salama 2010; Toma & 2010 Toma & Grigore 2010 Toma & Grigore 2010 Toma & Grigore 2014 Cambi & Cuadra Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. alternifolia L. Reaumuria germanica germanica 29 bracteata 40 Desv. Myricaria Tamaricaceae112 triandra ( thymifolia difolia ( salina pulverulenta pauciflora theses) paren- in onym (syn- species and Genus reuteri gran- ) ) 553 5 25 3 0 14 POWO species of Number eHALOPH In Species glands with Species ? 33 tutr Number/mm structure Gland + Weiglin 8 6+2 Olesen 8 6+2 6Gupta Gupta 56 62 cells of No CaSO 8 2 fflxN Cl / Na efflux and secreted elements Main CaCO SO Ca, K, Na, 4 4 3 notes & Reference 2010 Toma & Grigore 2010 Toma & ore Grig- 2014; al. et Cuadra 1989; Andres 2017 al. et D¨orken 1984 Murty & 1984 Murty & 2010 Toma & ore Grig- 1970; Smet De Denaeyer- 1991 Winter & 1979 Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. aphylla aphylla aphylla androssowii amplexicaulis albiflonum africana africana L Tamarix trigyna songarica negevensis hirtella theses) paren- in onym (syn- species and Genus . POWO species of Number 22 23 29 72 eHALOPH In Species glands with Species ? ? 34 tutr Number/mm structure Gland asdZhang Na, Cl, 8 Na, 6+2 Cl, 8 Raised 6+2 Raised 3 3Gupta Abbruzzese 23 43, 26 cells of No 2 fflxN Cl / Na efflux and secreted elements Main a K Na, Ca, Mg, Zn Na, Mn, Mg, K, Fe, Ca, Al, nmol/g/s SO Mg Ca K, Na, Cl, SO Mg Ca K, Na, Cl, SO Mg Ca K, SO Mg Ca K, 4 4 4 4 l8 Cl notes & Reference 1969 al. et Thomson 2017 al. et Santos 2008 Wang & Xue 1998 Ramadan 1972 Waisel 1984; Murty & 2013 al. et 1999 al. et Salama 1999 al. et Salama 2018 al. et Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. elongata dioica chinensis canariensis boveana austromongolica aucheriana arceuthoides arborea aphylla aphylla aphylla theses) paren- in onym (syn- species and Genus POWO species of Number eHALOPH In Species glands with Species ? ? ? ? 35 tutr Number/mm structure Gland uknZhang Zhang Zhang Zhang Sunken Sunken Raised Raised 8Gupta 48 cells of No Thomson 8 2 fflxN Cl / Na efflux and secreted elements Main Mg and K Ca, Cl, Na, Mo and Zn, Cu, Mn, B, SO4 and HCO3, NO3, Cl, Ca, Mg, K, Na, notes & Reference 1988 Waisel & Hagemeyer 1970 Berry 1967 Liu & 1984 Murty & 2018 al et 2018 al. et 2018 al. et 2018 al. et Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. meyeri mascatensis leptostachya laxa korolkowi karelinii jordanis indica hohenackeri hispida gracilis gracilis ganuensis gallica theses) paren- in onym (syn- species and Genus POWO species of Number eHALOPH In Species glands with Species ? ? ? 36 tutr Number/mm structure Gland uknZhang Zhang Zhang Zhang Zhang Zhang Sunken Raised Raised Zhang Sunken Raised sunken or Raised Sunken Ca, K, Sunken 8 36 Sessile 2Waisel Gupta 22 60 cells of No 2 fflxN Cl / Na efflux and secreted elements Main n Na and Mg notes & Reference 1972 1984 Murty & 1970; Smet De Denaeyer- 1993; al. et Bastos 2013; al. et Abbruzzese 2018 al. et 2018 al. et 2018 al. et 2018 al. et 2018 al. et 2018 al. et Zhang 2018 al. et 2018 al. et Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. smyrnensis ramosissima ramosissima ramosissima ramosissima ramosissima polystachya passerinoides parviflora nilotica nilotica theses) paren- in onym (syn- species and Genus POWO species of Number eHALOPH In Species glands with Species ? 37 tutr Number/mm structure Gland asdZhang Salama Salama 8 Raised 8 6+2 6+2 5Gupta 25 Gupta Gupta 12 24 cells of No 2 fflxN Cl / Na efflux and secreted elements Main 1974 lace Wal- and Kleinkopf Ba and Sr Mo, Ti, Si, Al, B, Mn, Cu, Ca, K, Na and Mn Mg, K, Ca, K and Na notes & Reference 2012 manaka Ya- & Acharya Imada, 2011 al. et Ma 1984 Murty & 1984 Murty & 1984 Murty & 2018 al. et 1999 al. et 1999 al. et Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. aet ta i 2A.docx Fig secretory-structures-in-the-plumbaginaceae-origin-evolution-and-roles-in-stress- al et Caperta file Hosted usneoides troupii tetragyna tarimensis taklamakensis smyrnensis theses) paren- in onym (syn- species and Genus POWO species of Number eHALOPH In Species vial at available glands with Species 9Waisel 19 ? https://authorea.com/users/302954/articles/433019- 38 tutr Number/mm structure Gland asdZhang Zhang Raised Raised 7Gupta 37 cells of No + Na, 6+2 2 fflxN Cl / Na efflux and secreted elements Main a bManousaki, Pb Ca, Ca K Cl, S, Al, Mg, notes & Reference 2008 akis Kaloger- & tonakis padan- Pa- Kadukova, 1972 2017 bye Weirs- & cock My- Wilson, 1984 Murty & 2018 al. et 2018 al. et Posted on Authorea 10 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158386324.48291028 | This a preprint and has not been peer reviewed. Data may be preliminary. tolerance 5.docx Fig secretory-structures-in-the-plumbaginaceae-origin-evolution-and-roles-in-stress- al et Caperta file Hosted tolerance 4.docx Fig secretory-structures-in-the-plumbaginaceae-origin-evolution-and-roles-in-stress- al et Caperta file Hosted tolerance 2B.docx Fig secretory-structures-in-the-plumbaginaceae-origin-evolution-and-roles-in-stress- al et Caperta file Hosted tolerance vial at available at available vial at available https://authorea.com/users/302954/articles/433019- https://authorea.com/users/302954/articles/433019- https://authorea.com/users/302954/articles/433019- 39