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Blumea 64, 2019: 92–95www.ingentaconnect.com/content/nhn/blumea https://doi.org/10.3767/blumea.2019.64.01.03RESEARCH ARTICLE

INTRODUCTION

In the course of the preparation of a manuscript on Boragina­ceae for the Flore du Gabon, some species of Heliotropium in the broad sense of Gabon and other African countries have beenstudied.Recently,Hilger&Diane(2003)rearrangedthegenericdelimi-tationandclassificationofBoraginaceaesubfam.Heliotropioi­deae(bysome,e.g.,Luebertetal.2016,recognisedatfamilylevel),basedonmoleculardataofnuclearITS1andchloroplasttrnLUAAintronsequences,combinedwithmorphologicalcha- racters.TheyproposedtomergeSchleideniaEndl.andHelio­tropium L. sect.OrthostachysR.Br. intoEuplocaNutt., andresolved MyriopusSmallasitssister.Alaterstudy,basedonmore markers and outgroups but only two Euploca species, indeed resolved these two genera as sisters, that are, together with Ixorhea Fenzl, forming the sister clade of the remainder of the subfamily being Heliotropiums.str.(Weigendetal.2014).EuplocawaspublishedbyNuttall (1836)toaccommodateanewNorthAmericanspecies(Euploca convolvulaceaNutt.)andwasuntilthemergerbyHilger&Diane(2003)anexclusivelyNewWorldtaxon.With the inclusion ofHeliotropium sect.Orthostachys and Schleidenia, Euploca now is considered to have a pantropi-caldistributionandtocontainc.100species(Luebertetal.2011),althoughcurrently inEuploca only68specificnamesexist(IPNI2018).However, a considerable number of recombinations has not yetbeenmade.Intheirstudy,Hilger&Diane(2003)notonlytransfer only a small selection of species, they also only make a single new recombination for a subspecies, while other existingsubspeciesandvarietiesof thespecies they trans-ferred,werenotdiscussednortransferred.AmongstthoseOldWorldspecimenstheyusedfortheirmolecularstudywasan

accession from Zambia, Gilges 685(M),ofHeliotropium baclei DC.var.rostratumI.M.Johnst.(asSchleidenia but that com-binationisnotvalidlypublished).Basedonthephylogeneticposition of this accession, they indeed created Euploca baclei (DC.)Diane&Hilgerbutdidnotmakeanynewcombinationforthetaxontheiraccessionbelongedto,var.rostratum.Thedifferences between typical Euploca bacleiandmaterialofvar.rostratumseemsolargethatwewonderedifthistaxonshouldnotberecognisedatspecieslevel.Although our research is focused on continental tropical Africa, westumbleduponacollection(J.M. Hildebrandt 3035)fromMadagascarofwhichtheKduplicate,anisolectotypeofEvol­vulus madagascariensisVatke,wasidentifiedasbelongingtoH.bacleivar.rostratum.Evolvulus madagascariensis is an older name than H. ka­tangense, theoldestavailablenameatspecies level forvar.rostratum, and hence the identity of this material is relevant for thenameofthecontinentalAfricanmaterial.So,weincludedmaterial of Heliotropium madagascariense(Vatke)I.M.Johnst.,asitiscurrentlyclassified,intoourstudies,resultinginacom-plete review of the African Euploca species with a widened and elongated beak, the Euploca bacleicomplex.

METHODS

HerbariumspecimensfromBR,BRLU,LandWAG,andtheMadagascarmaterialofMOhavebeenphysicallyexamined,whilespecimensfromE,KandPtogetherwiththosepresenton JSTOR-GlobalPlants (2017)were examinedusing highresolutionimagesontheinternet.AllexaminedspecimenswereenteredintheNaturalisdatabaseandgeoreferenced.Forsomemeasurementsalsotheimageshavebeenused.Formeasure-ments the beak length is measured from the sinus in the fruit-outlinetotheapexbutexcludingthestyleandstigmacapifthatisstillpresentandforfruitlengththebeakwasincludedaswell.

The Euploca baclei complex (Boraginaceae subfam. Heliotropioideae)E.L.A.N.Simons1,J.J.Wieringa1

1 NaturalisBiodiversityCenter, sectionBotany,P.O.Box9517, 2300RALeiden,TheNetherlands;

correspondingauthore-mail:erik.simons1978@gmail.com.

Key words

AfricabiogeographyBoraginaceaeendozoochoryEuplocaHeliotropiaceaeHeliotropiumkeymap

AbstractIn order to recognise both taxa previously regarded as varieties ofHeliotropium baclei, nowadays classifiedinEuploca,anewcombinationisnecessary.Asthetwovarietiesareclearlyseparableintermsofmor-phology and biogeography, we propose to raise these varieties to the species level, for which the new combination Euploca katangensisneedstobecreated.Moreover,weproposethenewcombination Euploca madagascariensis for Heliotropium madagascariense,aspeciesfromMadagascarconsideredbysomeasconspecificwithH. baclei, buttreatedhereasdistinct.Forthesethreespecieswithbeakedfruits,constitutingthe‘Euploca bacleicomplex’,akeyandadistributionmap,basedonrevisedherbariumspecimens,isgiven.Twoadditionalcombinations,Euploca bullockii and Euploca sessilistigma are made to complete the transfer of tropical African Heliotropium species that belong in Euploca.

Published on18March2019

93E.L.A.N.Simons&J.J.Wieringa:TheEuploca bacleicomplex

RESULTS

In our opinion, the two varieties of Heliotropium baclei are clearlyseparablebasedontwocharacters;lengthoftherostrumontheripefruitsandpetalcolour,asmentionedbyJohnston(1930),Heine(1963)andTaton(1971).Usingherbariumvouch-ers from the African continent we evaluated the mentioned morphologicalcharacters.Weconsiderthesedifferencestobequitesubstantial.Thelongbeaksandcompletelyyellowflowersofvar.rostratum are a striking difference with typical E. baclei.Moreover,typicalE. baclei has in general smaller fruits than the other variety, and there is even a gap in the ranges of the beakproportionoftheentirefruit length.ApartfromanareaalongtheNigerRiverinwesternMaliwheretheyco-occur,thetwotaxaaregeographicallyseparated,asisshowninMap1.We thereforewant to recogniseHeliotropium baclei var. ro­stratum at the species level, for which a new combination is proposed: Euploca katangensis.Interestingly,thematerialweexaminedofH. madagascariense seems more closely related to theWest-AfricanE. baclei, than to the pan-African E. katangen­sis.WithE. baclei it shares white flowers with a yellow centre, andrelativelyshortbeaks.FruitsofH. madagascariense are in general slightly smaller than those of E. baclei, but their width is slightly larger, resulting in a different length/width ratio with only a slight overlap: H. madagascariense fruits are wider than long(ratio0.6–0.95)whereE. baclei is about globose to longer thanwide(ratio0.9–1.8).Togetherwiththewidegeographicalgap between these two entities, we cannot accept this material asbelongingtothesamespecies.Hence,wefollowJohnston(1930)whoconsideredthisasaseparatespecies:Heliotropium madagascariense.AsHildebrandt 3035andotherMalagasyspecimensseenbyusclearly fit into the former subsectionAxillariaI.M.Johnst.underthesectionOrthostachysR.Br.andwe consider it closely related to E. baclei, we think it should be placed in Euploca as well, and hereby we propose a new combination.Sinceintheprotologuethereisnospecimendes-ignated as holotype for the type gathering, and the most likely one, theBsheet,mentionedasholotypebyFörther (1998),whichshouldbeconsideredasalectotypification,islost,weselecttheLsheetfromtheextantisolectotypes.

NOTES ON ECOLOGY, MORPHOLOGY AND KEY TO THE ‘EUPLOCA BACLEI GROUP’

Euploca can clearly be separated from Heliotropiums.str.butthecharactersarenoteasilyrecognisedinthefield.Euploca is characterised by mericarpid or endocarpid structures with surfacesculpturingsdescribedas‘pits’,kranz-chlorenchymaorganisationinleavesofalmostallspeciesandtheexclusiveoccurrence of characteristic trichomes on a pedestal of distinctly enlargedfoliarepidermiscells(Hilger&Diane2003).No comprehensive treatment of African Heliotropium has been writtenyet.However,followingJohnston(1928,1930),Förther(1998)andHilger&Diane(2003),mostAfricanspeciesnowa-daysclassified inEuploca formerly belonged to subsections Axillaria and Bracteata I.M.Johnst. of sectionOrthostachys.TheonlyexceptionisE. ovalifolia(Forssk.)Diane&Hilgerthatbelonged to subsection EbracteataI.M.Johnst.Speciesbelong-ing to former subsection Bracteataarequiteeasilyrecognisedby its bracteate solitary inflorescence and its pointed corolla lobes.SpeciesbelongingtoformersubsectionAxillaria have flowersbornealongaleafystem,notaggregatedintoadefinitespikeorracemeandhavedrupaceous,beakedfruits.Thus,apart from E. ovalifolia, African species of Euploca are easily distinctive from ‘true’Heliotropium s.str., like thepantropicalspecies H. indicumL.anditstypespeciesH.europaeumL.that have ebracteate, leafless, dichotomous spiciform inflo-rescences.Euploca ovalifolia has ebracteate spiciform, often dichotomous inflorescences too, but differs in some micromor-phological characters, typical for Euploca;havingaone-layeredendocarpid with surface sculpturings and having trichomes of theleavesonapedestalofenlargedepidermalcells(Förther1998,Hilger&Diane2003).Euploca bacleis.str.isasub-Saharan,UpperGuineantaxon,probablynotoccurringeastofMali (Map1).Euploca katan­gensisiswidespreadinAfricasouthoftheSaharaandeastofeasternMali.Euploca madagascariensisisendemictoMada-gascar.Ourmapdoesnotpresentallherbariumspecimens.Taton (1971)mentionssomecollections fromCentralAfricathat have not been seen by us, as is the case for Gilges 685 (M)fromZambiausedinthestudybyHilger&Diane(2003).It is also likelymorematerial fromMadagascarwill exist inherbariawedidnotvisit.The three taxaoccuratmuddy,sandyorgravellyshoresoflakes and river beds and other temporarily inundated areas, sometimesinroadsideverges,ditches,orevenontracks.Eu­ploca bacleihasbeencollectedinricepaddiesseveraltimes.Rees(1978)mentionsH.bacleivar.rostratum co-dominating drylagoonbottomsoftheKafueriverinZambiaduringthedryseason, together with Cynodon dactylon(L.)Pers.Theyseemto be pioneer species of open or almost absent vegetation, be-ingabletoquicklycovershoresthathaverecentlyfallendryorevenwhiletheyarestillinundated.InGabon,E.katangensis wascollected(Lachenaud 2024)atthestonyshoresofalakein the sedimentary basin of theOgooué river.At least thetypespeciesofthegenus(E.convolvulacea)seemstohaveasimilarecology(Nuttall1836).Dispersalofthefruitscouldpossibly occur by means of endozoochory as is the case with manyplantspeciesinaquatichabitats(Soonsetal.2016).Andindeed, fruits of E.katangensis have been found in the guts of birds,atleastinknob-billedgeese(Sarkidiornis melanotos)aninter-tropicalmigrantspecies(Douthwaite1978).Euploca bacleioccursinWest-Africanplainsof0–200mabovesea level, E.katangensis is frequent in south- andeasternAfrican highlands at about 1 000 m elevation, reaching up to 1 690 m in Ethiopia, but in Gabon it has been collected at ele-vationsofonly0–20m.MostrecordswesawofE. madagas­

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Map 1 Distribution of Euploca katangensis(GürkeexDeWild.)E.L.A.N.Simons&Wieringa(),E. baclei (DC.)Diane&Hilger()andE. mada­gascariensis (Vatke)E.L.A.N.Simons&Wieringa().

94 Blumea – Volume 64 / 1, 2019

cariensis come from the lowland but one record from an area atc.800melevation.All three species grow decumbent to ascendant, often starting inrosettes.Theleavesareabitcoriaceous,darkgreenandabitglossy,coveredwithwhiteappressedhairsonbothsides.Thecorollaistrumpet-shaped,4–7mmlong,thedryfruitsarebeaked.DeCandolle(1845)doesnotmentionwhetherH.baclei isaperennialoranannual;neitherdoesDeWildeman(1903)for H.katangense,whileTaton(1971)explicitlyreferstoH.ba­cleivar.rostratumasperennial.Indeedsomeoftheplantsarebecomingquitewoodyatbase.Inonlyfourofthespecimensexaminedbyusthehabitwasexplicitlymentioned:J. Chillou 1357 and J.V.G. do Espirito Santo 2005 regarded E. baclei as annual, as is the case with collections of E. katangensis(Drum­mond 6264, 6762)fromZambia.Wethinkbothspeciescouldbeannualorperennial,dependingonecologicalcircumstances.

Key to the species in the Euploca baclei complex

1. Flowersyellowtoorange-yellow.Fruitswithlong(1.6–4.2mm)beak,longerthanremainderoffruit:beak/fruitlengthratio0.5–0.7 . . . . . . . . . . . . . . . . . . . . . . . E. katangensis

1. Flowerswhitewith a yellow centre. Fruitswith a shorter(0.2–1.1mm)beak,shorterthanremainderoffruit:beak/fruitlengthratio0.2–0.4 . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Fruitswiderthanlong,length/widthratio0.6–0.95.—Mada-gascar . . . . . . . . . . . . . . . . . . . . . . . .E. madagascariensis

2. Fruitsaboutaswideas longor longer, length/widthratio0.9–1.8.—WestAfrica . . . . . . . . . . . . . . . . . . . . E. baclei

CONSERVATION ASSESSMENT

Sincebothcontinentalspeciesaccordingtoourdataoccuratover 10 locations over a fair range, and without imminent threats weconsider both species as LeastConcern (LC).Euploca madagascariensis seems more restricted with fewer locations and might be endangered, but since we have only seen a part of the available material, we refrain from making a full assess-mentforthisspecies.

NOMENCLATURE AND NEW COMBINATIONS

Euploca katangensis(GürkeexDeWild.)E.L.A.N.Simons&Wieringa,comb. nov.—Fig.1;Map1

Heliotropium katangenseGürke exDeWild.,Ann.Mus.CongoBelge,Bot.sér.4, [1(3)] (1903)223.—Heliotropium bacleiDC.var. rostratum I.M.Johnst.(1930)91.—Type:E. Verdick 141(BR)(lecto,designatedbyFörther1998),Congo(Kinshasa),Katanga,Lukafu,Oct.1899.

Heliotropium nigerinumA.Chev. (1920)45.—Syntypes:A.J.B. Chevalier 1168(P),Mali,Sebi,11July1899,A.J.B. Chevalier 1365(P),Mali,surlesbordsdumarigotdeDay,3Aug.1899.[Note:oftenconsideredasnomen nudum, but since Chevalier gives a description, we consider this name validlypublished.]

Heliotropium marifoliumauct.nonRetz:Baker&Wright(1905)40.

Note—TheauthorshipofH. katangenseisratherunclear.ThenameispublishedwithassoleauthorGürke,butinapaperwrittenbyDeWildeman,wheretheintroduction(atpagevii)thanksseveralworkersinBerlin,includingGürke,fortheirhelp,butonlyofK.Schumannitisexplicitlymentionedhemadethedescriptionsforhisnewspecies.WethereforehavetoassumeonlythenamewascoinedbyGürkeandthedescriptionisbyDeWildeman.

Fig. 1 Euploca katangensis(GürkeexDeWild.)E.L.A.N.Simons&WieringainZimbabweZambeziRiver,ZambeziNationalPark18-01-2016.—Photo:BartWursten.

95E.L.A.N.Simons&J.J.Wieringa:TheEuploca bacleicomplex

Euploca madagascariensis(Vatke)E.L.A.N.Simons& Wieringa,comb. nov.—Map1

Evolvulus madagascariensisVatke,Linnaea43(7)(1882)522.—Heliotro­pium madagascariense(Vatke)I.M.Johnst.—Type:J.M. Hildebrandt 3035 (lectoL,designatedhere,barcodeL0004002;isoBM,BREM,GH,GOET,JE,K,LE,M,P,W).

Asstatedintheintroduction,manytaxa,especiallyfromSouthAmerica, that belong in Euplocahavenotyetbeentransferred.SincewehavenoopiniononthestatusofmanynamesfromtheNewWorldwewill not dealwith those, but for tropical Africathereareonlytwoacceptedtaxathathavenotyetbeencombined in Euploca, so by making these combinations at least thispartoftheworldwillhavebeendealtwith.Thereforeweproposethenexttwocombinations:

Euploca bullockii(Verdc.)E.L.A.N.Simons&Wieringa,comb. nov.

Basionym.Heliotropium bullockii Verdc., Fl.Trop.E.Africa,Boragin.(1991)75.—Type:A.A. Bullock 2284(holoK).

Euploca sessilistigma (Hutch.&E.A.Bruce)E.L.A.N.Simons&Wieringa,comb. nov.

Basionym.Heliotropium sessilistigmaHutch.&E.A.Bruce,Bull.Misc.Inform.Kew1941(2)(1942)160.—Type:J.B. Gillett 4107(holoK).

AcknowledgementsWethankBartWurstenforhispermissiontorepro-ducethephotographofFig.1,andwethanktwoanonymousreviewersforreviewingthemanuscript.WethankthecuratorofMissouriBotanicalGardenforsendingustheirspecimensfromMadagascar.’

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IDENTIFICATION LIST

Collectornames(initialsonlyaddedwhenrelevant forAfrica)andcollec-tion numbers, followed by the herbarium at which the specimen is present (acronyms followingThiers 2017).Collectionsmarkedwith * havebeeninvestigatedbasedondigitalspecimensonly(MNHN2016,JSTOR-GlobalPlants2017,RoyalBotanicalGardenEdinburghHerbariumCatalogue2017).Recordsmarkedwith**arereliablerecordswithphotomaterialinobserva-tionorphotodatabases,butwithnoexsiccatespecimens.AllrecordshavebeengeoreferencedandenteredintheNaturalisdatabase.

Euploca baclei (DC.)Diane&HilgerAdam13646(WAG);14866*(P).Bacle*(G-DC,GH)–Berhaut1690(BR,P);4084*(P);7172*(P)–Breman268(WAG).

ChampluvierS101(BR,LG,WAG)–Chevalier1148*(P)–Chillou1357(IFAN,P,WAG)–A.Cissé135(WAG).

EspiritoSanto2005(BR,WAG);2055(LISJC,WAG);2474(LISJC,WAG).Hagerup309(BR).Lécard42(BR,WAG)–Lejoly87/10(BRLU)–Lisowski56428(BR).MortonSL1357(FHI,GC,IFAN,K,SL,WAG).J.Raynal7787*(P).C.VandenBerghen1966(BR);2041(BR);4292(BR,P);5339(BR,WAG)–Verdellens.n.Gambia1972(L.2747523).

Wailly5154*(P).

Euploca katangensis(GürkeexDeWild.)E.L.A.N.Simons&WieringaAchten634(BR)–Auquier4352(L,LG,WAG).Bidgood6060(BR)–Bingham10741(MRSC,SRGH,WAG)–A.A.Bullock1198(BR);2331(BR).

Chevalier1168*(P);1365*(P)–Christiaensen701(BR,WAG).DeSaeger101(BR)–J.Dekker**(N/A)–Drummond6264*(E);6762*(E).Fanshawe3887(BR)–Fotius1620*(P)–Friis2072(BR,WAG).R.G.A.Germain594(BR)–Gilges685(M).Hagerup317(BR)–Hooper1941*(P).Lachenaud 2024 (BRLU,G, LBV,MO,P,WAG) – Lisowski 27173 (BR,WAG);52012(BR).

Reekmans3754 (BR)–M.A.E.Richards7080 (BR);11758 (BR);20549(BR)–E.A.Robinson5434(BR).

M.Sanane1519(DSM,K,WAG)–Symoens6821(BR).Troupin246(BR);702(BR,WAG).Vanderyst4591(BR);4629(BR)–Verdcourt3447(BR)–Verdick141(BR);182(BR).

Wailly5078*(P)–Witte5271(BR)–Wursten**(N/A).

Euploca madagascariensis(Vatke)E.L.A.N.Simons&WieringaBardot-Vaucoulon1858*(P).J.M.Hildebrandt3035*(BM,BREM,GH,GOET,JE,K,L,LE,M,P,W)–Humbert2079(MO,P);4086(MO).

Rakoto1377(MO)–RakotovaoRN5785(MO)–Randrianaivo962(CNARP,MO,P,TAN).