Division Haptophyta – phylogeny and diversity

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  1. 1. Division Haptophyta phylogeny and diversity Bente Edvardsen, Elianne Sirns Egge & Wenche Eikrem Ecologyanddiversity Haptophytesaremostlysingle-cellednanophytoplankton(usually 2-30m)presentinallseas.Someoccurinfreshwater.Theyareone ofthemajorgroupsofprimaryproducersintheocean,andarealso importantgrazersonpicoplanktoninopen-ocean.Some haptophytes,suchasmembersofEmiliania,Gephyrocapsa, Phaeocys1s,ChrysochromulinaandPrymnesiummayformextensive bloomsthatmayaecttheglobalcarbonbalanceandclimate forcing,orcauseshkillswithecologicalandeconomicalimpact. Duetotheirsmallsize,speciesarediculttodisKnguishbyopKcal microscopy,andknowledgeontheirdiversityanddistribuKonis incomplete.Molecularmethods,suchasenvironmentalsequencing andmetabarcoding,hasrevealedalargediversityofhaptophytesin thesea,manywithinthepicoplanktonsizefracKon.Mostphylotypes donothaveclosesimilaritytoculturedandsequencedspecies,and someformcladesconsisKngofenvironmentalsequencesonly,that mayrepresentnovellineagesattheclasstogenuslevel(Fig.1). EdvardsenB,EikremW,GreenJ,AndersenRA,Moon-vanderStaaySY&MedlinLK,2000:PhylogeneKcreconstrucKonsoftheHaptophytainferredfrom18SribosomalDNAsequencesandavailablemorphologicaldata.Phycologia39:1935. EdvardsenB,EikremW,ThrondsenJ,SezAG,ProbertI&MedlinLK,2011:RibosomalDNAphylogeniesandamorphologicalrevisionprovidethebasisforarevisedtaxonomyofthePrymnesiales(Haptophyta).Eur.J.Phycol.46:20228. EggeES.,EikremW&EdvardsenB2015a:Deep-BranchingNovelLineagesandHighDiversityofHaptophytesintheSkagerrak(Norway)Uncoveredby454pyrosequencing.-J.Eukaryot.Microbiol.62:12140. EggeES,JohannessenTV,AndersenT,EikremW,BihnerL,LarsenA,SandaaR-A&EdvardsenB,2015b:SeasonaldiversityanddynamicsofhaptophytesintheSkagerrak,Norway,exploredbyhigh-throughputsequencing.Mol.Ecol.inpress. ThrondsenJ,HasleGR,andTangenK,2007.PhytoplanktonofNorwegianCoastalWaters,AlmaterForlagAS,Oslo. Fig. 1. Maximum likelihood (RAxML) tree based on haptophyte 18S rDNA sequences of all cultured species and of environmental clones (284 taxa). The tree includes all major taxonomic groups and deep-branching clades without a cultured representative, potentially representing novel classes (HAP-2-5) or orders (clade D, E). Modified from Egge et al. 2015b. Fig. 2. Drawings of haptophytes:(a)Haptolinahirta(Prymnesiaceae, Prymnesiales),(b)Chrysochromulinaalifera(Chrysochromulinaceae, Prymnesiales),(c)Phaeocys1spouche1i(Phaeocystales),(d)Coccolithus pelagicus(Coccolithales),and(e)Pavlovagyrans(Pavlovales).FromThrondsen etal.(2007). 1 0 Department of Biosciences, University of Oslo, Norway bente.edvardsen@ibv.uio.no +CladeEandF ? ? chloroplast haptonema flagella scales Fig3. *Drawingof Prymnesiumparvum byJahnThrondsen. *ThinsecKonof Haptolinafragariacell wheren=nucleus, m=mitochondrion, c=chloroplast, f=agella,p=pyrenoid. TEMmicrographby WencheEikrem. *Scalesoftwolife cyclestagesof Prymnesiumpolylepis. TEMmicrographby BenteEdvardsen. *SEMmicrographof thecoccolithophorid Emilianiahuxleyiby EysteinPaasche. *LMmicrographof Prymnesiumparvum cellsbyCanter-Lund& Lund. Taxonomy,morphologyandlifecycle Haptophytapresentlycomprisesapprox.320species,80genera, 5-7orders(Pavlovales,Phaeocystales,Prymnesiales, Coccolithales,Isochrysidales,(Zygodiscalesand Syracosphaerales))and2classes(Pavlovophyceaeand Prymnesiophyceae).ItismonophyleKcandproposedtobelong tothesupergroupHacrobia.Mostareunicellular,planktonic agellates,butcoccoid,amoeboid,palmelloid,lamentous, colonialandbenthicformsalsooccur(Fig.2).Mostare phototrophicwith1-4goldenbrownchloroplasts(Fig.3),but someareheterotrophicandmanyaremixotrophic.Thetypically havetwosmoothagellaandathirduniqueappendage,the haptonema,thatcanbelongandcurling,toshortandsKor vesKgial(Fig.3).Itcanbeusedtoahachtoasurfaceorforfood handling.Cellsaretypicallycoveredbyorganicscales,and coccolithophoridcellsalsobycalciedscales,coccoliths(Fig.3). SpeciesidenKcaKonislargelybasedonscalemorphology observedbyelectronmicroscopy.Haptophytesmayhavea heteromorphichaplo-diploidlifecycle. References: 0.08 HQ877921_Isochrysidales_Isochrysidaceae_Pseudoisochrysis_paradoxa_Lee_et_al_2011_CULT AM491021_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_simplex_Edvardsen_et_al_2012_CULT KC888101_Isochrysidales_Isochrysidaceae_Chrysotila_lamellosa_Bendif_et_al_2014_CULT AY919672_Kathablepharis_remigera HM581604_Prymnesiophyceae_X_Prymnesiophyceae_XX_Prymnesiophyceae_XXX_sp_Cuvelier_et_al_2010_ENV JX680352_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_sp_Simon_et_al_2013_ENV AJ544121_Coccolithales_Pleurochrysidaceae_Pleurochrysis_dentata_Saez_et_al_2003_CULT KJ763246_Prymnesiophyceae_X_Braarudosphaeraceae_Braarudosphaeraceae_Lie_et_al_2014_ENV JN938582_Isochrysidales_Isochrysidaceae_Isochrysis_sp_Liu_et_al_2012_CULT JX660987_Phaeocystales_Phaeocystaceae_Phaeocystis_globosa_Decelle_et_al_2012_CULT JX680402_Prymnesiales_Prymnesiaceae_Prymnesium_polylepis_Simon_et_al_2013_ENV AM490995_Prymnesiales_Prymnesiaceae_Haptolina_brevifila_Medlin_et_al_2012_CULT HQ869379_Prymnesiales_Prymnesiaceae_Prymnesium_sp_Orsi_et_al_2012_ENV KF899845_Prymnesiales_Prymnesiaceae_Dicrateria_rotunda_Gilg_and_Preston_2014_CULT HQ394158_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Domaizon_et_al_2012_ENV KC888109_Isochrysidales_Isochrysidaceae_Isochrysis_galbana_Bendif_et_al_2014_CULT JX291802_Prymnesiales_Prymnesiales_Prymnesiales_Thompson_et_al_2012_ENV L04957_Isochrysidales_Noelaerhabdaceae_Emiliania_huxleyi_Bhattacharya_et_al_2001_CULT HQ868752_Clade-F_Clade-F_X_Clade-F_XX_sp_strain1_Orsi_et_al_2012_ENV HM581609_Prymnesiales_Prymnesiaceae__Cuvelier_et_al_2010_ENV HM103447_Isochrysidales_Noelaerhabdaceae_Emiliania_Gephyrocapsa_sp_Quaiser_et_al_2011_ENV GU370008_Prymnesiales_Prymnesiales_X_Prymnesiales_XX_sp_Lin_et_al_2009_ENV HM135063_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_parva_Luo_et_al_2011_ENV KC888120_Isochrysidales_Isochrysidaceae_Isochrysis_galbana_Bendif_et_al_2014_CULT JX680418_Isochrysidales_Isochrysidales_X_Isochrysidales_XX_sp_Simon_et_al_2013_ENV JF714228_Pavlovales_Pavlovaceae_Exanthemachrysis_sp_Bendif_et_al_2011_CULT AJ246276_Isochrysidales_Noelaerhabdaceae_Gephyrocapsa_oceanica_Edvardsen_et_al_2001_CULT KC888119_Isochrysidales_Isochrysidaceae_Isochrysis_galbana_Bendif_et_al_2014_CULT JX660986_Phaeocystales_Phaeocystaceae_Phaeocystis_globosa_Decelle_et_al_2012_CULT EU500065_Prymnesiophyceae_X_Prymnesiophyceae_XX_Prymnesiophyceae_XXX_sp_Frias-Lopez_et_al_2009_ENV JF714230_Pavlovales_Pavlovaceae_Pavlova_sp_Bendif_et_al_2011_CULT JX680404_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Simon_et_al_2013_ENV EU500073_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Frias-Lopez_et_al_2009_ENV HQ865173_Phaeocystales_Phaeocystaceae_Phaeocystis_pouchetii_Orsi_et_al_2012_ENV AM491019_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_parva_Medlin_et_al_2012_CULT KC404141_Isochrysidales_Noelaerhabdaceae_Emiliania_huxleyi_Bendif_et_al_2013_CULT FJ431495_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_sp_strain35_Marie_et_al_2010_ENV HQ869040_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_sp_strain35_Orsi_et_al_2012_ENV FJ431487_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Marie_et_al_2010_ENV FJ537310_Coccolithales_Coccolithales_X_Coccolithales_XX_sp_Shi_et_al_2009_ENV FR874498_Prymnesiales_Prymnesiaceae_Imantonia_sp_Newbold_et_al_2012_ENV KC488451_Prymnesiales_Prymnesiaceae_Haptolina_sp_Dasilva_et_al_2013_ENV EU500076_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Frias-Lopez_et_al_2009_ENV AF107082_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Moon-van_der_Staay_et_al_2000_ENV AM779755_Prymnesiales_Prymnesiaceae_Prymnesium_palpebrale_Seoane_et_al_2012_CULT KF130129_Prymnesiophyceae_X_Braarudosphaeraceae_Braarudosphaeraceae_Wu_et_al_2014_ENV AJ246268_Prymnesiales_Prymnesiaceae_Prymnesium_nemamethecum_Edvardsen_et_al_2002_CULT JX660995_Phaeocystales_Phaeocystaceae_Phaeocystis_sp_Decelle_et_al_2012_CULT JX680354_Pavlovales_Pavlovaceae_Diacronema_vlkianum_Simon_et_al_2013_ENV HQ394114_Prymnesiales_Prymnesiaceae_Prymnesiaceae_X_sp_Domaizon_et_al_2012_ENV HQ877902_Isochrysidales_Isochrysidaceae_Isochrysis_galbana_Lee_et_al_2011_CULT AM491026_Coccolithales_Calcidiscaceae_Oolithotus_fragilis_Medlin_et_al_2012_CULT JF714226_Pavlovales_Pavlovaceae_Exanthemachrysis_sp_Bendif_et_al_2011_CULT JF714243_Pavlovales_Pavlovaceae_Diacronema_noctivaga_Bendif_et_al_2011_CULT KC404153_Isochrysidales_Noelaerhabdaceae_Gephyrocapsa_oceanica_Bendif_et_al_2013_CULT HQ867320_Clade-HAP-5_X_Clade-HAP-5_XX_Clade-HAP-5_XXX_sp_Orsi_et_al_2012_ENV AB183588_Pavlovales_Pavlovaceae_Pavlova_sp_Sekiguchi_et_al_2004_CULT JX680435_Phaeocystales_Phaeocystaceae_Phaeocystis_antarctica_Simon_et_al_2013_ENV AF107083_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_X_sp_Moon-van_der_Staay_et_al_2000_ENV JX680370_Phaeocystales_Phaeocystaceae_Phaeocystis_antarctica_Simon_et_al_2013_ENV FJ537342_Clade-HAP-2_X_Clade-HAP-2_XX_Clade-HAP-2_XXX_sp_Shi_et_al_2009_ENV HM474582_Phaeocystales_Phaeocystaceae_Phaeocystis_sp_Lepere_et_al_2011_ENV JF714223_Pavlovales_Pavlovaceae_Exanthemachrysis_gayraliae_Bendif_et_al_2011_CULT KF129696_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_sp_Wu_et_al_2014_ENV KC888102_Isochrysidales_Isochrysidaceae_Chrysotila_sp_Bendif_et_al_2014_CULT KF129833_Prymnesiales_Chrysochromulinaceae_Chrysochromulina_sp_Wu_et_al_2014_ENV AM490984_Zygodiscales_Pontosphaeraceae_Scyphosphaera_apsteinii_Medlin_et_al_2012_CULT AB183616_Coccolithales_Pleurochrysidaceae_Pleurochrysis_sp_Sekiguchi_et_al_2004_CULT HM474577_Phaeocystales_Phaeocystaceae_Phaeocystis_sp_Lepere_et_al_2011_ENV KC404124_Isochrysidales_Noelaerhabdaceae_Emiliania_huxleyi_Bendif_et_al_2013_CULT FJ157339_Prymnesiales_Prymnesiaceae_Prymnesium_sp_Genitsaris_et_al_2009_ENV HM565914_Prymnesiales_Clade-B3_Clade-B3_X_sp_Cuvelier_et_al_2010_ENV DQ207406_Pavlovales_Pavlovaceae_Diacronema_noctivaga_Herve_et_al_2005_CULT JN693092_Prymnesiales_Clade