introduction of six novel species from morphology, phylogenetics, and toxin production, withthe An evaluation of the genus Amphidinium (Dinophyceae) combiningevidence Harmful Algae

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Research Paper

An evaluation of the genus Amphidinium (Dinophyceae) combining evidence from morphology, phylogenetics, and toxin production, with the introduction of six novel species

Sarah Karafas

^a,

*, Sing Tung Teng

^b

, Chui Pin Leaw

^c

, Catharina Alves-de-Souza

^d

aAlgalResourcesCollection,UniversityofNorthCarolinaWilmington,MarineBiotechnologyFacility,5600MarvinK.MossLn.,Wilmington,NC28403,USA

bFacultyofResourceScienceandTechnology,UniversitiMalaysiaSarawak,94300KotaSamarahan,Sarawak,Malaysia

cBachokMarineResearchStation,InstituteofOceanandEarthSciences,UniversityofMalaya,Bachok,16310Kelantan,Malaysia

dAlgalResourcesCollection,UniversityofNorthCarolinaWilmington,MarineBiotechnologyFacility,5600MarvinK.MossLn.,Wilmington,NC28403,USA

ARTICLE INFO

Articlehistory:

Received4May2017

Receivedinrevisedform1August2017 Accepted1August2017

Availableonline29August2017

ThismanuscriptisdedicatedtoDr.Theo- doreJ.Smayda(1932–2017)inhonorofhis lifetime dedication to phytoplankton ecologyandserviceasformereditorof HarmfulAlgae.

Keywords:

Secondarystructure Compensatorybasechange Cytotoxicity

Scales 28S ITS

ABSTRACT

ThegenusAmphidiniumisanimportantgroupofathecateddinoﬂagellatesbecauseofitshighabundance inmarinehabitats,itsmember’sabilitytoliveinavarietyofenvironmentalconditionsandabilityto producetoxins.Furthermore,thegenusisofparticularinterestinthebiotechnologyﬁeldforitspotential inthepharmaceuticalarena.Taxonomicallythethereisahistoryofcomplicationandconfusionoverthe properidentitiesandplacementsofAmphidiniumspeciesduetohighgeneticvariabilitycoupledwith highmorphologicalconservation.Thirteenyearshaspassedsincethemostrecentreviewofthegroup, andwhilemanyissueswereresolved,someremain.Thepresentstudyusedmicroscopy,phylogeneticsof the28SregionofrDNA,secondarystructureoftheITS2regionofrDNA,compensatorybasechangedata, andcytotoxicitydatafromAmphidiniumstrainscollectedworld-widetoelucidateremainingconfusion.

Thisholisticapproachusingmultiplelinesofevidenceresultedinamorecomprehensiveunderstanding ofthemorphological,ecological,andgeneticcharacteristicsthatareattributedtoorganismsbelongingto Amphidinium,includingsixnovelspecies:A.ﬁjiensis,A.magnum,A.paucianulatum,A.pseudomassartii,A.

theodori,andA.tomasii.

1.Introduction

ThegenusAmphidiniumisagroupofathecateddinoﬂagellates thatareincrediblydiverseinthat,whilebeinghighlyconserved morphologically, they thrive world-wide in a wide variety of habitats(Dodge, 1982;DolapsakisandEconomou-Amilli,2009;Flø Jørgensenetal.,2004a,2004b;Larsen,1985;LarsenandPatterson, 1990), temperatures(Murray and Patterson,2002), and trophic modes(FløJørgensenetal.,2004;Murrayetal.,2004;Murrayand Patterson, 2002).They also producetoxins and bioactivecom- poundsthatcanhavebothharmfuleffects.Reportsindicatethat Amphidinium blooms cause ﬁsh kills and that the toxins they producemayincreasetheeffects ofCiguateraFishPoisoningas theyareoftenfoundinassociationwithGamberdiscusinaffected

areas(Baigetal.,2006;Rhodesetal.,2010;TindallandMorton, 1998).Theyalsohavebeneﬁcialeffects,producingcompoundsthat exibitantifungalorantimicrobialproperties(Echigoyaetal.,2005;

Kobayashi andKubota,2007; Kobayashiet al.,1991; Kobayashi, 2008;Mengetal.,2010;Nuzzoet al.,2014;Satakeetal.,1991;

Washida,2006).Theygroweasilyinculture,andcanbescaledup tomassculturingvolumesappropriateforchemicalanalyses.

Presentlythegenusincludesbothheterotrophicandautotro- phicformspossessingacharacteristicallyminuteepiconethatis deﬂectedtowardstheleft.TheautotrophicAmphidiniumconsistof twoclades(FløJørgensenetal.,2004a,2004b)thataresistertoone another.TheﬁrstwillbereferredtoastheHerdmaniiCladeand includes:A.steiniiLemmerman,A.mootonorumMurrayandD.J.

Patterson,A.herdmaniiKofoidandSwezy,andA.cupulatisquamaM.

TamuraandT.Horiguchi.ThesecondistheOperculatumCladeand includes:A.carteraeHulburt,A.massartiiBiecheler,A.gibbossum(L.

Maranda and Y. Shimizu) Flø Jørgensen and Murray, A. trulla Shauna Murray, Rhodes, and Flø Jørgensen, A. operculatum Claparède & Lachmann, and, most recently, A. thermaeum

*correspondingauthor.

E-mailaddresses:[email protected](S.Karafas),[email protected] (S.T.Teng),[email protected](C.P.Leaw),[email protected]

(C.Alves-de-Souza).

HarmfulAlgae68(2017)128–151

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