ALLOTOONIA, A NEW NEOTROPICAL GENUS OF APOCYNACEAE BASED ON A SUBGENERIC SEGREGATE OF ECHITES

2004·9.3 MB·English

by  J Francisco Morales

ALLOTOONIA,ANEWNEOTROPICALGENUS
OFAPOCYNACEAEBASEDONA
SUBGENERICSEGREGATEOFECHITES
J.FranciscoMorales Justin K.Williams
InstitutoNacionaldeBiodiversidad(INBio) DepartmentofBiologicalSciences
Ap(o22-3100,SantoDomingodeHeredia Saml-loustonStateUniversity
COSTARICA Huntsville,Texas77341-2116,U.S.A.
ABSTRACT
AJloUwnia(Apocynaceae)anewneotropicalgenussegregatedfromEchilesisdescribed.Inaddition,
akeytospecies,descriptions,distributionmaps,illustrationsandspecimensexaminedareprovided
forallspecies.
KeyWords.Apocynaceae,Apocynoidcac,Al/oloonm.Echiteae,Echi(es,Neotropics,Mesoamerica,
Mexico
RHSUMEN
AUotoonia(Apocynaceae),unnuevogeneroneotropicalsegregadodeEchitesesdescrito.Sebrinda
unaclaveparatodoslostaxa,asicomodescripciones,mapasdedistribucion.ilustracionesy
especlmenesexaminadosparatodaslasespecies.
InhismonumentalrevisionofEchitesP.Browne,Woodson(1936)established
twosubgenera:Echites(Euechite.s)andsubg.PseucJechitesWoodson.Subgenus
PseudechitesincludedthespeciesEturbinataWoodsonandE.tuxt/ensisStandi.
andwasdistinguishedfromsubgenusEchitesbyitssmallercorollas(1.25-2.5
cm longvs.5-8cmlong),lobesnarrowlyoblongandreflexed(vs.obliquely
obovatcandspreading),anditsmanyflov/eredlaxinflorescence(vs.compact
andfewflowered).Athirdspecies,E.parvifloraSesse&Mog.wassubsequently
addedtosubg.PseudechitesbyWoodson(1938).
Echitesw^entrelativelyunnoticedforcloseto60years,untilMorales(1997)
providedasynopsisofthegenus.Inhiswork,Moralesdescribedanadditional
speciesinsubg.Pseudechites(E.puntarenensisj.F.Morales)aswellaspresented
anargumentformaintainingPrestoniaagglutinata(Jacq.)WoodsoninEchites,
resurrectingtheoldernameEchitesagglutinata.
ThemorphologicalcladisticanalysisofEchitess.l.andothergeneraofsub-
familyApocynoideaepresentedinWilliams(2004)indicatesthatEchitessensu
Woodsonispolyphyletic.SubgenusPseudechitesshowsastrongeraffinityto
othergeneraofEchiteaesensuEndressandBruyns(2000)(e.g.,Laubertia.^
Prestonia,Thenardia)thanitdoestosubg.Echites.Consequently,theauthors
havechosentorecognizethespeciesinsubg.Pseudechitesasadistinctgenus
(AUotooniadescribedbelow)separatefromEchitess.s.Table1documentsthe
SIDA21(1):133-158.2004
13^1 BRIT.ORG/SIDA21(1)
differencesbetweenAllotoonia,Echiies,andotherselectedgeneraofNewWorld
Echiteae.
MORPHOLOGICALCHARACTERS
—
Leaves. Theleavesareoppositeandentire.Fourofthefivespecieshaveleaves
withobscuresecondaryvenationmakingthemreadilydiscerniblefromEchites.
Onespecies, however,A.agglutinata,hasconspicuoussecondary veins.
AUotooniaalsoshareswithothersneotropicalApocynoideae,intra-petiolar
colleters.Allspecieshaveglabrousleaves.TheleavesofA.turhinataareoften
twotothreetimeslongerandbroaderthanthoseoltheothertourspecies.Field
studieshaveshownthatthesecharactersareuniformandconsistentwithina
population. —
Inflorescence. AllotooniaisreadilydistinguishedfromEchitesbythein-
florescencestructure.TheinflorescenceofAllotooniaisahelicoidcymeonce,
twicetooccasionallythricebranched,whileinEchitestheinflorescenceisa
modifieddichasium,sometimesappearingracemose,andoccasionallyreduced
tojustoneflowerThebractsarealwaysinconspicuousandscarious.
—
Flowers. Thecalyxisregular,pentamerous,andwithasolitaryopposite
colleterontheadaxialsurfaceofeachsepal,asfoundinmanyothersmembers
oftheEchiteae(e.g.,Prestonia,Thenardia).Thesepalsarenarrowlyovateand
T-3.5mmlong.Theshapeanddimensionsofthesepalsforeachspeciescorre-
spondstotheshapeanddimensionsofthebractssubtendingthepedicel.
Thecorollaissalverform,thetubeglabrouswithout.Theaestivationis
dextrorseasareallotherneotropicalmembersoftheApocynoideae,Theco-
rollalobesareverynarrowlyelliptictoalmostliliform,acuminatetolong-
acuminateattheapexandconspicuouslywavyandtwisteddistaWy.Allotoonia
istheonlyneotropicalApocynaceaegenuswithwavyandtwistedfiliformlobes
attheanthesis(Figs.1-2).InEchites,thecorollalobesareobovate.
Thestamensareincludedandtheanthersarestronglyattachedattwolev-
elstothespoolshapedstylehead(Fig.3).Theanthershapediffersbetween
AllotooniaandEchites(Fig,4).InAUoLooniaznantherismoreorlesscontinu-
ous(exceptforaslightdorsalprotuberance),withoutamarginalrib.InEchites
theantherhasaconspicuousmarginalribthatisextendedfromthebasalpro-
jectionsal—ongtheentirelengthuptotheapex(Fig.1B).
Fruit. ThefruitsofAllotooniaarecomposedoftwoherbaceous,pendu-
lous,follicles(Fig.5)developingfromtwocarpelspostgenitallyunitedatthe
apex,sometimesfreeatmaturity.Thefolliclesarecontinuoustoobscurely
moniliform.ThefruitsofEchitesarewoodyandnevermoniliform.
Seeds.—TheseedsofAIlotooniaareglabrous,6-20perfollicle,androstrate,
withthebeakslender,narrowlyandconspicuouslyelongate(Fig.6),whilein
Echitestheseedsareinconspicuoustomoderatelyrostrate,withthebeakshort
MORALESANDWILLIAMS,ALLOTOONIA,ANEWNEOTROPICALGENUSOFAPOCYNACEAE 135
(sometimessomewhatelongateinE.yucatanensis)andgraduallyacuminate
towardthemicropylarend(Fig.7).
AUotooJn.Fi.aMJo.rFa.leMsor&alJ.eKs.W&iJl.lKi.amWsilliams,gen.nov.Type.AUotooniaagglutinala(Jacq.)
Herbavolubilis,Forsteronieae,Laubertiae,Parsoniae,Prestonieae.ThenardiaeetThorcauiaeaffmis:
foliaoppositapetiolata,membranacea,eglandulifera;inflorescentialateralispedunculatamultiflora;
corollaregulans5-partita,hypocrateriforma,lobisliliformis,revolutis,tuboexappendiculatastamina
inserta,glabra;fructusfolliculis,seminibusrostrata.
Lactescentlianas,usuallyherbaceous,morerarelysuffruticose,notwoody.
Stemsteretetosubterete,somewhatflattenedatthenodes,glabrous,not
lenticellate,withinconspicuousintrapetiolarcolleters,thelatexusuallywa-
teryLeavesopposite,entire,membranaceous,glabrousorglabrate,thesecond-
aryandtertiaryveinsusuallyinconspicuousorobscure,petiolewithmany
minuteandfiliformcolletersmtheaxils.Inflorescenceahehcoidcyme,axil-
lary,glabrous,many-flowered,usuallylong-pedunculate,bractsscarious,in-
conspicuous.Flowerspentamerous,thesepalsfreeorveryslightlyimbricate
basally,scarious,bearingasolitary,episepalousentiretovariouslylaciniate
coUeterwithin;corollasalverform,glabroustoveryminutelyandinconspicu-
ouslypuberulentwithout,withoutcoronalobesorannularcorona,thetube
usuallylongitudinallygrooved,withaninfrastaminalribspresentbeloweach
filament,thelimb5-parted,actinomorphic,dextrorselyconvolute,thelobes
verynarrowlyelliptictoalmostfiliform,acuminate,twistedandwavydistally;
stamens5,included,insertedintheupperpartofthecorollatube nearthe
mouth,anthersconniventandagglutinatedtothestylehead,attachedintwo
pointstothestylehead,glabroustoglabrate,thecaewiththebasesagittate,2-
auriculate,auriclesshort,butconspicuous,short-acummate,filamentsshort,
puberulenttopilose,carpels2,unitedattheapexbyacommonstylarshaft
surmountedbythefusiform,spool-shapedstylehead,ovaryglabrous,ovules
numerous,multi-seriate,nectarinesfive,separatedanddistinct,rarelyslightly
connatebasally.Folliclescontinuousandteretetosubterete,orobscurely
moniliform,slenderandsmooth,usuallyglabrous;seedscomoseatthe
micropilarend,conspicuouslyrostrate,minutelyrugose,numerous.
ThisgenuscomprisesfivespeciesdistributedfromsouthernMexicoto
northernPanama,andtheWestIndies.AllotooniaisnamedafterDrAnthony
Leeuwenberg,fromtheWageningenUniversity(WAG),TheNetherlands,in
recognitionofhisworkmtheApocynaceae.
ThegenusAllotooniaisinthesubfamilyApocynoideae,tribeEchiteae(En-
dress&Bruyns2000),asevidencedbythefollowingcharacters:antherswith
thethecaeconnivent,agglutinatedandstronglyattachedattwolevelstothe
spoolshapedstylehead.
Table1,MorphologicalcharactersinAliotooniaandseieciedgeneraufiheEchiteuc
Genera Aliotoonia Echites Forsteronia Laubertia Prestonia Thenardia Thoreauea
Morphological
Characters
Latex Clear While White Clear White Clear White
Secondaryveins Usually Conspicuous Conspicuous, Conspicuous Conspicuous Usually Conspicuous
inconspicuous rarely inconspicuous
Inconspicuous
Inflorescence Helicoidcyme Modified Thyrslform Scorpioidcyme, Racemose, Subumbellate Trichotomously
dichaslum sometimes subracemose, cyme branched
reducedand umbellate. Subumbellate
appearingsimple subumbellate, cyme
orumbelliform corymboseor
subcorymbose
Bracts Scarious Scarious Scarious Scarious Foliaceousto Scarious Scarious
scarious
Sepals Scarlous Scarious Scarious Scarious Foliaceousto Scarious Scarious
scarious
Sepals Solitaryand Solitaryand Laterallyorevenly Lacking Solitaryand Solitaryand Solitaryand
colleters opposite opposite disposed,rarely opposite opposite opposite
acking
Corolla Salverform nfundibuliform Salverformtorotate Salverform Salverformto Rotate Urceolate
rarely tocampanulate
infundibuliform oo
Corona Absent Absent Absent Present, Present, Absent Present, o
continuous continuous dissectedinterior S!
around around tocorollamouth 3>
thecorolla thecorolla NJ
mouth mouth
so
Genera Allotoonia Echites Forsteronia Laubertia Prestonia Thenardia Thoreauea m
Morphological O3>
Characters
5
Lobes Vfeilellriiyfpotrincma,rtoroawllmyost Obovate Olabnlcoenoglatteo Obovate Oovbaotveateto Ovate Ovate yiooz:r»->
acuminate,
twistedandwavy m
distally
Stamens Included, Included, ncludedor Includedor Includedor Exserted Apicallyexserted Oo
insertion insertedinthe insertedabout variously variously apically 2
tuhpepemroupatrht,near cmoirdolwlaaytutbhee exserted exserted exserted 3am>t
Filaments Free Free Freeorconnate Free cFroenenaotre Partially Partiallyconnate o-3n>
Diskor 5,usually 5,butusually Usually(3-)5- 5,separatedto Annular,entireto 5,separated 5,separated
nectaries separated,rarely connate lobed irregularly 5-iobedor
slightlyconnate basally basally connatein5 divided m
basally nectaries
Follicles Slender, Slender, Slender, Slender, Divaricate, Slender, Unknown
continuousand continuous divaricateor, moniliform,rarely fusiform. moniliform
terete,or andterete parallel. continuous continuousor
obscurely continuous rarelymoniliform
moniliform tomoniliform
Comaseeds Conspicuously Inconspicuously Truncate,sessile Truncate,sessile Truncate Sessile Unknown
rostrate to
moderately
rostrate
K^
13!5 BRIT.ORG/SiDA21(1)
f\G.^.AIIotooniacaudata.A.Habitandinflorescences{Morales8242,INB).B.Close-upoftheflowers,showingtwisted
andelongatecorollalobes{Morales8242,INB).
KEYTOTHESPECIESOFALLOTOONIA
1. Corollalobes(17-)20-27mmlong;corollatubeconstrictedbelowtheinsertionof
thestamens;montanerainforestandrelateddisturbedareasbetween1500-
2350m A.turbinata
1. Corollalobes4-14(-16)mmlong;corollatubestraightorslightlyconstrictedbe-
lowtheinsertionofthestamens;tropicaldeciduousforest,premontanemoistfor-
est,galleryforest,andrelateddisturbedareas,bewteen0-1200(-1400)m.
2. Secondaryvenationofleavesraisedandconspicuous,tertiaryvenationofleaves
obscure;corollalobesglabrous;plantsoftheWestIndies A.agglutinata
2. Secondaryvenationinconspicuous,notraised,tertiaryvenationusuallyincon-
spicuous;corollalobespubescentorglabrous;plantsofMexicoandCentral
America.
3. Comromlladilaomb.e,scsohnosrttreicrttehdanbteutbwee,egnlambirdodulse;btuodbpaosientofedl;otbuebse.7-9mmlong,ca.A.tuxtlensis
3. Cmomrollloangl,obcea.s2lomnmgerditahma.n,stturbaei,ghhti,rsnuottecoornsgtlraibcrtoeuds.;budround;tube4-6(-7)
4. Corollaorangetoyellow-orange,thelobeshirsute;pedicels2-9mm;an-
thers3.5-4mm A.parvifiora
4 Corollawhite,creamishwhitetopinkishwhite,thelobesglabrousto
glabrate;pedicels11-24mm;anthers(5-)6-8mm A.caudata
MORALESANDWILLIAMS,ALLOTOONIA,ANEWNEOTROPICALGENUSOFAPOCYNACEAE 139
fi6.2.Allotooniaparviflora.h.Habitandinflorescences(Morales&Abarca88W,INB).B.Close-upoftheflowers,show-
ingtwistedandelongatecorollalobes[Morales&Abarca8810,INB).
1.Allotooniaagglutinata(JacqJJ.F.Morales&J.K.Williams,comb.nov.(Figs.8,
9). Basionym:EchitesdgglutinataJacq.,Hnum,Syst.Pi.13.1760.Anechites
agglutinata(Jacq.)Miers,Apocyn.S.Amer.236.1878.Prestomaagglutinata(Jacq.)
Woodson,Ann.MissouriBot,Gard.18:552.1931.Type.HAITI:Cap.Francais,Select
Stirp.Am.Hist.1763.(lectotype,heredesignated).
EchilescircinalisSw.,Prodr.52.1788.Hacmadictyonciixiimlis(Sw.)G.Don,Gen.Hist.4:83.18.37.
Ar\echilescircinalis(Sw.)Miers,Apocyn,S.Amer.236,1878,TYPE:datalacking(fl),Rohr93
(lectotype,heredesignated:C;ISOLECTOTYPE:P-LAj,
EchitessanguinokntaTussac,Fl.Antill.95,t,11,Haemadictyonnutans(Anders,)A.DC.van
sangmnolenta(Tussac)A.DC,Prodr8:426,1844,TYPE.HISPANIOLA:notlocated,
EchitescircmahsSw,varthomasianaA.DC.,Prodr466,1844,Anechitesihomasiana(A.DC.)
Miers,Apocyn.S.Amer237.1878,Type:STTHOMAS:exactlocalitylacking,1841(fl),
Friedrichsthal240(holotype:W),
EchitesohtusijoliaSesse&Moc,Naturaleza(MexicoCity),ser2,2,app.45,1893.Type:PUERTO
RICO:nearCastellumdelMorro,Oct..yearlacking,Se.s.se&Mocino5075(LECTC^TYPE.here
designated:MA).
Suffruticosetoherbaceousliana,stemteretetosubterete,glabroustoglabrate.
Leaves:petioles5-20mmlong,withminutepectinatecoUetersintheaxilsor
sometimeseglandular,glabrous;blade3-13.5x2-8.5cm,membranaceous,el-
140 BRIT.ORG/SIDA21(1)
^l ^^-K ^
I fei
irm
: '^^m \'\
',«•--'/.;: :
k
A B -ivj
mm mm mm
2 1.5 3
Fig.3./l//ofoo/i/flparviflorastyleheadandanthers,show- Fig.4.AllotooniaandEchitesanthers,dorsalview.A.
ingthetwolevelsofattachmenttothespoolshapedstyle Allotooniaparviflora.(Morales3139,INB).B.Echites
head.{Morales&Abarca88W,INB). umbellata.{Whiteford8209,INB).
liptic,narrowlyelliptictonarrowlyovate,apexcaudatetoobtuse,baseobtuse
tosomewhatattenuatebasally,glabrous,midveinimpressedonbothsurtaces,
thesecondaryvenationconspicuousonbothsides,hillorescenceusuallylonger
thanthesubtendingleaves,la.x,peduncleb5-7cm,pedicels2-5mm,bracts1-2
mmlong,ovate,acute,scarious;sepals1-L5x0.5-1mm,ovate,acuminate,gla-
brous,glabratc,orminutelyandinconspicuouslypuberulent,colletersome-
whatlaciniatetodeeplyso;corollasalverform,white,creamishwhitetocream,
glabrouswithout,tube5-6mmlong,1-1.5mmdiam,somewhatinllatedatthe
attachmentol thestamens,pubescentwithinnearthemouth,lobes4-6x1-2
mm,glabrous;stamens:lilamentsca.4mm,glabratetosparselypubescent,
anthers4-5mm,glabrous,auriclesca.0.2mm,ovaryca.2mm,glabrous,style
head1.5-2mm long,nectaries1-1.5mm,entireandconspicuouslyseparate.
Follicles11-31C-46)x 0.4-0.8cm,continuous,smoothandglabrous,usually
straight;seeds6.5-10x1.5-2mm,rugose,thetannishcoma2-4cm.
MORALESANDWILLIAMS,ALLOTOONIA,ANEWNEOTROPICALGENUSOFAPOCYNACEAE 141
Fig.5./l/Zofoon/opaAi'/flora.Follicles
142 BRIT.0R6/SIDA21(1)
2cm
hQ.e.Allotooniaseeis.H,A.agglutinataiAxelrod&Axelrod3091,\M).B.A.turbinata{Gomez-Letal.,14142,m).C.A.
tuxtlensis{Arvigoetal.536,m).D.A.pamflora(Morales3161,INB).
cm cm
1 1
Fid.7.ffWfesseeds.A.E.umbellata{Trejoetal.530,INB).B.E.yucatanensii(Magallanes3137,INB).