Table Of ContentSTUDIESINNEWWORLDAMARANTHUS
(AMARANTHACEAE)
JAMESHENRICKSON
DepartmentofBiology
CaliforniaStateUniversity
LosAngeles,CA 90032. U.S.A.
ABSTRACT
Anutrcinthuscrassipesisrecognizedasconsistingoftwovarieties:var.cn/ss/pcsfromtiie
WestIndies,coastalMexico-Texas-Louisiana,andnorthernSouthAmerica,andvar.uYiruockii
oftlieCliiluiahuanDesertregionofTexasandiVIexico.Anuvcintbusscleropoicle.scontainstwo
morpiisthatarenottaxonomicallyrecognized.Aiiic/ranthnsXtexensis,consideredofhybrid
originbetweenA.cnissipesandA.scleropoides,isdescribedasnew.Amaranthmberlandieriis
notconsideredasdistinctfromthemorewidetin^K^^A.polygomncks.Anewspecies,Amamnth/is
tamaidipe>nh,isdescribedfromsouthmostTexasandeasternMexico.ItisrelatedtoA.
dubiiiiandthepoorlyknownA.hrandegeiofSinaioa.AnenigmaticAmaranthnsxtucsonetish
isdescribedfromtheTucsonMountainsofsouthernArizona,withrelatedpopulationsoccurring
inSonora,Mexico.
RESUMEN
SereconoceaAnuircinthnscrassipescomoconstituidapordosvariedades:lavar.crassipes,
condistribuciondesdelacostadeMexico,TexasyLouisiana,atravesdelCaribehastael
nortcdcSudamenca;ylavar.warnockii,delaregiondelDesiertoChihuahuense.Amaranthiis
scleropoidesconsistededosfasesmorfologicassindesignacionderangotaxonomicoformal.
Lanoroespecienut\c\Amaranthusxtexensisprobablementeseoriginodelahibridacionen-
treA.crassipesyA.scleropoides.NoseconsideraaA.berlandiericomodiferentedeA.polygonoides,
especiedemasamj-)liadistribucion.SedescribeAmaranthustamaultpensis,especienueva
delestedeMexicoylapartemasmeridionaldeTexas.EstaespecieestarelacionadaconA.
duhiusyconA.hrandegei,taxonsinaloensepobrementeconocido.Sedescribelaenigmatica
AmaranthusXtucsonensiscomonotoespecienuevadelaSierradeTucson,delsurdeArizona.
KkywoRixs;Amaranthus,Amaranthaccae,planttaxonomy,plantsystematics,hybridization.
StudiesintheAmaranthaceaeinconnectionwiththeChihuahuanDesert
FlorahaveresultedinthispaperonthesystematicsoftheAmaranthuscrassipes-
A.scleropoidesandA.polygonoidesgroups.Twoadditionaltaxaaredescribed
asnew.
TheAmaranthuscrassipes-A.scleropoidesgrouphasdichasialclustersofflowers
borneatallnodesfromthebaseoftheplanttothetip.Theinflorescences
aremodifiedcompounddichasiainwhichtheterminalflowersareovertopped
byopposite,sinuouspeduncles thatbearonetoafewpairsofpersistent
bractswithmostlymaleflowersandterminateinpistillateflowersthatare
againovertopped bysimilarpaired,sinuouspeduncles.Asthepeduncles
SiDA18(3):783-807.1999
—
784 SiDA 18(3)
maturetheybecomegreatlythickenedand induratedinsomespeciesand
trapthefruitbetweentheexpandedpeduncles(Fig. 3A,D).Gray(1862)
consideredthickenedpedunclestobeanabnormalcharacterinAirass/pes,
butHolzinger(1892)showedtheywereconsistentlyproducedinthatspe-
cies,causedbyaproliferationofexpandedparenchymacellsthatdevelop
thickenedwalls.Atmaturity,theentireinflorescencefallsfrom theplant
andservesasadispersalunitcapableofflotation.
VARIATIONWITHINAMARA.NTlllLS(.RASSIPES
AmaranthuscrassipesSchltdl.isawidespreadNewWorldspeciesocctir-
ringfromtheWestIndies,Bahamas,FloridaKeys,Arizona,Texas,Mexico,
Colombia,andVenezuela(Fig. 1) itstypelocalityisonSt.Thomasinthe
VirginIslands.Thespeciesconsistsofdecumbent-procumbent,glabrous,
monoeciousannualswithovate-obovatetolanceolate,long-petiolateleaves.
Theflowersareproducedateachnodeusuallyfrom thebaseoftheplant
upwards inshortdichasia 5—10 mm long (Fig. 2C, E). Thedichasiade-
veloptheaforementionedgreatlythickened,indurate,stronglysinuouslateral
axes(Fig.3A).Thefemaleflowershave5erecttospreading,narrowlyspattilate
sepalsthatarethickenedandjoinedatthebase(Fig.3B).Theyareusually
shorterthanthefruit.Thefruitwalliscloselyadherenttotheseedandindehiscent,
althoughitmaydevelopacoloredencirclinglinebelowthemiddle.Itis
usuallysmoothbelowandsomewhattuberculateaboveandalongthemar-
gins.The2(—3)stylesarejoinedtogetherintoadistinctivelyrate-shaped
neckbeforetheytaperanddivergeasslendertips(Fig. 2A—C).Themale
flowersareproducedonthesidesofthedevelopinginflorescenceaxes.They
matureandfallawaybeforetheinflorescenceaxesthickenandtheyhave5
membranoussepalsandthreeanthers.
ThroughouttheCaribbeanregiontheplantsareinitiallyerectbutusu-
allybecomeprocumbentweedsindisturbedplaces.Theyusuallyhaveovate,
broadlyovatetoobovateleavesthatmayeitherbeuniformlysmallor,in
fastgrowingplants,largeinsize.(Fig2C).Similarovate-leavedplantsalso
occurinnorthernSouthAmerica,theYucatanPeninsula,northeastMexico,
coastalTexas,Louisiana,KeyWest(Florida)aswellasSonoraandArizona.
Incontrast,plants in inlandTexasand in theC^hihuahuanDesertregion,
developnarrower,moreoblanceolate,grayishleaves(Fig.2E).Theyalsotend
tobe moreerectordectmibent ingrowth habit, features thatareshared
withtherelatedA.scleropoides.Thegrayishcoloroftheleavesalsomakethe
reticulatepatternoftheKranzstructurelessconspicuous.In1944I.M.Johnston
describedtheseinlandplantsasAmaranthuswaniockii,distinguishingthe
speciesbyitselongate,somewhatthinneroblanceolateleaves,itslesselon-
gate,moreslenderstems,andsmallermorecompactcymesconspicuously
crowdedatthestembases.Exceptforthenarrower,moreoblanceolate-lan-
Henrickson,StudiesinNewWorldAmaranthus 785
Fig. 1.DistibiicionotAmamnthuscraisipesvar.crassipesandvar.wcirnockii.
ceolateleaves,alltheotherfeaturesnotedbyJohnstonalsooccurinocca-
sionalplantsof/\.crassipesthroughoutitsrange.Inalltechnicalcharacters
offlowers,fruitandinflorescence,A.warnockiiandA.crassipesareidentical.
Recognitionofthesetwotaxaatthespecieslevelisconsideredunjustified
as thedistinguishingfeaturesareminorand tosomeextent inconsistent
andrecognitionwouldcreatepoorlydifferentiatedmicrospecieswithinthis
highlyvariablegenus. ButoverallthegroupofspecimensreferabletoA.
warnockiicanberecognizedsolelyonthebasisofthenarrower,moreoblan-
ceolate,grayishleaves.BecauseofthisA. warnockiiishereretainedatthe
varietallevelasitrepresentsageographicallybasedsubunitofthespecies
(Fig. 1).CorrellandJohnston(1970),incontrast,consideredA. warnockii
synonymouswithA. crassipes.ThecollectionsfromsouthernArizonaand
adjacentSonora,however,havedistinctlyovateleavesandfallwithinvar.
crassipes.AformaldescriptionofA.crassipesfollows.
AMARANTHUSCRASSIPES
Lowspreading,taprooted,procumbenttodecumbent,basallybranched,
monoeciousannuals,sometimeswiththecentralstemserect-ascending;stems
glabrous,striate,stramineous,moderatelybranched, 1—4(—9)dmlong;in-
ternodes 1.5—4cmlongbelow,reducedto5—15mmlongdistally.Leaves
alternate,ascending,(12—)30—55(—78)mmlong,thepetiolesslender,(5—)
10-25(—47)mmlong;leafbladesbroadlytonarrowlyobovatetoovate,to
1
786 SiDA18(3)
obhmceolateorelliptical,(7-)2()-30(^3())mmlong,{5-)l()-2()(-28)mm
wide,obtusetorounded,usuallyretusewiththemidveinshort-excurrent
at thetip, narrowlyacuminate-cuneateatthebase,themarginsentireto
erose-crisped,whitish,theveinscurvedtowardsthetip,butnotextending
tothemargins,distinctlywhitenedbeneath,thelaminaglabrous,withdistinct
Kranzinternalanatomy.Flowersborneinaxillary,congested,tanorrarely
dark brown,glomeratedichasia5—9mmlongusually borneatall nodes
fromthebaseoftheplantupwards,thedichasiadecussatelybranchingbelow
eachpistillateflower,thebranchesstronglysinuate-curved,becomingthickened
avenldopiinndgurfarutietd;,br0a.c9t—sa.n2d(—br1a.c6t)eomlemsbtrhoiacdklyandedltcatreo,wd0i.n5g—-0e.9ncmlomsilnogngt,hgeredeen-
alongthemidrib,themarginsscarious,broadlyattachedatthebase,per-
sistingonthesidesofthethickeneddichasia.Staminateflowersproduced
onthesidesofthedevelopingdichasia,with5,subequal,lanceolatesepals
1.2-1.6mmlong,thesemembranousexceptwheregreenalongthemid-
rib,acute;stamens3;anthers0.7—0.9mmlong;filaments1.5—2mmlong.
Pistillateflowersbornateachbranchofthedichasium;fruitingsepals5,
with2ateachmarginofthefruitandwithasmallersepalononeface,2on
ltohnego,thtehrefcalcea,wsthneasrerpoawlslydilsitnienacrt,ly0.c1l-a0w.e2domrmmerweidley,spkaeteulleadt,e,ex1p.4a—n2d.i3ngmmto
ovate,acute,slightlykeeled,conduplicate,sometimesslightlyreflexedtips
0.5—1 mmlongandnearlyaswide,thetipseachwithagreeninatruUate
patchmediallyandwithmembranousmargins,thesepalsexpandingtobecome
spongy-indurate,tan,unitedandtruncatedaroundthefruitbase.Fruiting
ovarycompressedorbicular-obovate, 1.3-1-5 mmwideandlong(exclud-
ingthestyle),thefruitwallsomewhatthickened,closelysurroundingthe
seed,indehiscentorinsomeplantswithadarkenedlineinthelowerhalf,
usuallyburnotconsistentlysmoothbelow,distinctlytuberculateaboveand
alongthemargins,toppedwiththepersistentstyle;thestyles2(—3),about
1.5mmlong,connateanderectfor0.2—0.5mmbelow,theyellowishlobes
erectorusuallylyrate,curvingoutward,theninwardastheytaper,termi-
natingasslenderoutwardlycurvedappendages,thepapillate-stigmasex-
tendingallalongtheinnersurfaceofthestylesontotheterminalslender
p0o.r6t—i0o.n7s.mSmeedtshiccokm,psrheisnsneyd,-doabrokvobirdo,wn1-.b1—la1c.k3,mtmhelmoanrg,gi0n.9o-b1t-us1emtmoswoimdee-,
whatrounded,thesurfaceshiny,withavisiblecellularsurfacepattern.With
twovarieties:
AmaranthuscrassipesSchltdl.var.crassipes(Figs.2A—C,2A—B).Linnaea
6:757.1831(as/\wrfm/////,0.5'r/tT«/wjmm/y7ef(Sthltdl.)Moq.inDC.Prod.13-^:271.
WI(ES4S9;TE/IixNoDltIiEiScn.issSitp.esT(hSochmlatsdl.I)slHainedr.on(.A,dBorli.vuAlcoasd.inNsaulca.eCSot.rdTohboam4a:e1,3.118<2SH6I-.1T8y2pi8;):
:
Henrickson,StudiesinNewWorldAmarantiius 787
Fic;.2.Fruitsandfoliai^eolAwayanthiiscrussipes.A-C;.A.crassz/'csvar.cras.u/KS.A.JVlattire
fruit:notecharacteristiclyratestylesandclawedsepalsthatareunitetlatthebase(Cameron
Co.,Tex.,Rnnyoii6yH,TEX).B.JVIatureFruit(Anti^aia,Box.57),NY).C.Characteristic
ovateleafblades(Apalachicola,Fla.,C/^z/w/^/ffHerk33^3ANY).T)-E.A.m/.u/pcsvar.icaninckii.
D.MatureIruit(PresidioCo.Tex.,Hinckley129,SRSC).E.Characteristicoblanceolateleaf
bB,laDde,s=(T2ercremllfCoor.C,;,TEe.x.m,nmk&Johnston17209,SRSC).WhitebarinA - 1 mmforA,
Ehrenk'i-gs.n.(holotyph:HAL).Dr.U.Braunhassentaxeroxcopyoftheholotype
fromHAL,depositedTEX.
Sciero[)t/sciuuivdnthouksSchrad.,Indexsem.Hort.^'ott. 1835;(reprintedin:Linnaea11
Litt.-ber.89. 18.t7).Typh:unknown.
Plantstypicallyerectwhenyounj^'butbecomingprocumbent,occasion-
allydecumbent,leafbladesovatetobroadlyovate,sometimesnearlyorbicularor
obovate,(7-)l5-30{-50)mmlong,(5-)10-l5(-28)mmwide,thelowersurface
greentoyellow-green,distinctlymarkedbythewhitish,arcuateveins.
Weed inroadsides,wastegrounds, fields,pastures,inclay,silty,some-
timessandyflats,beaches,androckyslopesinshadedoropenhabitats,widesjiread
intheCaribbeanregion,southFlorida,coastalLouisiana,Alabama(adven-
tive),Texas,northeasternMexicosouthtonorthernSouthAmerica(Columbia,
Peru),fromsealevelto25()(-]250)m(Fig. 1).ItalsooccursinArizona,
Sonora,Chihuahuaathigherelevations.Collectedthroughouttheyearbut
788 SiDA18(3)
expectedtobevegetativelyactiveduringhotmonthsduetoC4photosyn-
theticpathway.
AmaranthuscrassipesSchltdLvanwarnockii(I.M.Johnston)Henrickson,
cArobmorb..2n5o:v1.53(.Fig19.421.D-TEyp)i.-:BMasEiXoInyCmO:.Am/Cno/afiju/fuji/.zas:wca/nm/oitk/S/E].oMf.OJcohanmsptou,m,l)o.wArpnloalcde
nearmot,'oteonplain,8Sep19-11,l.M.JohustaiiH8ti6(n()].(ri-YPn:GH!).
Plantsinitiallyerectwhenyoung,thelateralbranchesbecomingdecumbent,
a3s3c(e-n4d4i)ngmtmerlmoinnga,ll(y4;-l)e6a-flb(l)ad(e-sl6o)blamncmeolwaitdeet,ontahrersouwrlfyacoebslamnocreeolgaltaeu,c(ou1s2--g)re1e6n-^
theveinsofthelowersurfaceslender,notconspicuous.
Opensikyfiats,mesquitethickets,muddyareasto1300melevation,in
south-westTexasand intheChihuahuanDesert inCoahuilaand eastern
Chihuahua(Fig. 1).
AA\ARAN'rnUSSCLI-ROI'OIDES
In 1895UlineandBraydescribedasecondspeciesrelatedtoA.crassipes
differmginitscircumscissile,smoothutricleand 3stylebranches. Uline
andBrayconsideredAmaranthusscleropoiclesintermediatebetweenA.crassipes
andA.graecizansautb.)!onL.(nowA,alb//sL.),thelatteralsohaving3styles
andcircumscissileutricles.Vegetativelyandininflorescencestructure,A.
scleropoiclesisverysimilartoA.crassipes.butA.scleropoiclesisataller,bushier
plant.Whileitsmatureutriclesareuniformlycircumscissile,thepistillate
flowersaresomewhatvariable insepal structure,stylenumberandother
features.Maturesepalsvaryfrom1.l-l."^mmintotallength.AsinA,crassipes.
twosepalsareusuallydistributedateachedgeofthematurefruit,witha
twoononefaceand aonesmallersepalon theotherface.Thesepalsare
clawedandvarygreatlyindevelopment.Insomespecimensthebladepor-
tioniserect,green,rounded,apiculateandratherflattened,however,inothers,
includingthetypecollection,thetipsaredistinctlythickened,keeled,con-
duplicate,recurved,andsharppointed,withacentralgreenrhomboidpatch
(Fig.4C).Stylesvaryfromtwotothree(Fig. 1A-B),orrarelyfive(I-flower
onCory2933GFI),withsomeplantsbeingconsistentlytwostyledandothers
withvaryingnumbers.Inflowerswithtwostyles,thestylesarelocateddirectly
overthemarginsofthecompressedovaryandasthefruitmaturethestyles
developdistinctdecurrentridgesthatextenddownthefrukmargins.When
stylesarethree,thethird,oftenreducedstyle,developsononeofthefaces,
againwithadistinctdecurrentbase.Thesurfacesoftheutriclemaybesmooth
ortightlyturberculed.
Amongcollectionsobserved,twodistinctformsarerecognizablebased
ondifferencesintliedevelopmentofthestylesandupperovarywall.The
mostcommonformhasdistinctlyspongy-thickenedstylebasesandtheupper
,
Hi-XRiCKSON,StudiesinNewWorldAmaranthus 789
Fk;. 3.AmaranthuscrassipesandA.sderopoides.A-B.A.crassipesvar.crassipes. A.Mature
dichasiaiinflorescenceshavespongy-thickened,induratedlateralbranches;theexternalbracts
subtendedmaleflowers,theFemaleflowersarelocatedatthebaseandripsoftheinflores-
cencebranches(Jamaica,Prodor23937,NY).B.Maturefruit-calyx(Jamaica,Proctor23957
NY).C-E.A.scleropoicks.C.Youngplantwithcharacteristicnarrowleafblades,axillary
inflorescences.D.Matureinflorescence.E.Maturefruitandcalyx(allWilliamsCo.,Tex.,
Wcilcott303,TEX).MagnificationsmA&BholdforD&E.
790 SiDA18(3)
utricle is induratedandstramineousincolor(Fig.4A-B).Thestylesare
papillate(stigmatic)alongtheirinnermarginsand thepapillaecontinue
ontotheslenderterminallobes.Asecondformdoesnotdevelopexpanded,
indurated thickeningson thestylebasesand upper h'uitand the twoor
threestylesarethusclusteredatthetopofthefr—uit(Fig.4C).Thetwoforms
bothoccurthroughouttherangeofthespecies theydonotsortoutgeo-
graphically(Fig. 5). Becauseofthis,and becausethisvariation does not
correlatewithanyothermorphologicalorecologicalvariation,theyarehere
recognizedasunnamedforms.Adescriptionofthespeciesfollows.
AmaranthusscleropoidesUline&Bray,Bot.Gaz 19:316. 1894.(Figs.
3C-E,4A-C,5).Tvi'i::U.S.A.TEXAS:"Tcxa.s,WesternTexastoElPaso,"WYii^ht
^tS'i;(i.ixTOTVPi;:CjH!;isoi.hc'i'oim'i;GH!).NotationonlectotypebyEM.Jolinston;
"Texas:ValVerdeCounty,Wrightfieldntimber7ycS',Amaranth,Devil'sRiver,stim-
mitofhills,2Aug1849."Thetypehassepalswithcondu|ilicate,sharplydivergent
tips;thestylebaseisnotinflatedasinsomecollections.
VerysimilartoAmarcinthuscvcissipesvegetativelyandininflorescenceand
flowerorganization,buttypicallymoreerect,bushy,2-5dmtallandoftenas
wide,withthelowerstemsascendingordeciunbentandascendingtoerect
mm
distally.Leaveswithslenderpetioles(4-)l()-25(-47) long;leafblades
mm
lanceolate,oblanceolatetonarrowlytrullate,(3^)15-25(-42) long,(2)
5-1()(-12)mmwide,glabrous,somewhatglaucous.Flowersborneinthickened,
sinuatedichasiaproducedfromthebasetothetopoftheplant;staminate
flowerswith5sepals,3stamens.Pistillatefruitingsepals5,with2ateach
marginofthefruit,twosepalsontheoneface,withonesmallestsepalon
theotherface,thesepalsmoderatelyclawedtonarrowlyspatulate,in(ruit
(1.2-)I.5-2.2(-2.3)mmlong,theclaws0.2-0.7mmwide,thedistalsepal
bladetypicallybroader,erectorvariouslyreflexed,±keeledandcondupli-
cate,±ovate-lanceolate,acute,withasmallorlarge,central,greennarrow
totrullategreenpatchto0.5-1 mm long,themarginsmembranous,the
sepalsexpandingandbecomingspongy-indurate,tan,andunitedaround
mm
thefruitbase. Fruitingovarycompressedorbicular-obovate, 1.1-1.3
wideandlong(excludingthestyles),thewallcircumscissillydehiscent,with
adistinctbrownishmedialline,thetxxlysmoothorbecomingcloselyturberculate
aboveandlessstronglytuberculatebelowthelineofdehiscence,thestyles
2-3(-5)tthethird(fifth)one(s)smallerwhenpresent},archingoutwardfrom
theuppermarginsofthefruitwall,thestylebasescontinuingasdecurrent
ridgesdownthefruitwall,thelowerportionotthestylesandadjacentdis-
talfruitwall(butnotextendingdowntothelineofdehiscence)oftenspongy-
thickened,inflated,orthedistalfriutwallandadjacentstylebasesonlyslightly
inflatedandascending;thestigmaspapillate,extendingalongtheinnermargin
ofthestylelobesandcontinuingasslendertipsfor0.4—0.5mm.Seedscompressed
obovoid, 1.0-1.1 mmlong,0.85-0.96mmwide,0.6-0.7mmthick,dark
Hknrickson,StudiesinNewWorldAmaranrhus 791
brown—black,themarginobtusetosomewhatrounded,thesurfaceshinny,
withaslightlyvisiblecellularsurfacepattern.
ANF.WTAXONFROMSOUTHERNTEXAS
Within thecomplexofA. crassipesandA. siieropoides. aseriesofspeci-
mensfromsouthernTexasarecharacterizedbyanerect-ascendinggrowth
habitandwhatappeartobecircumscissileutriclesasinA.scleropoides.However,
theutriclesarethickwalled,weaklyandcloselytuberculateandareactu-
allyindehiscentasinA.crassipesalthoughadistinctmediallineispresent.
AsinA.crassipes,thestylesareunitedatthebaseintoadistinctneckbefore
theyseparate,butthe2—3styleshavedistinctdecurrentridgesthatextend
downtheutriclewallas inA. scleropoides(Fig.4E).Thesespecimensalso
haverathernarrowlylanceolate tolinear-lanceolate, longpetioledleaves,
and theytend tohavelonger,morestronglyreflexedpistillatesepalsofa
typesometimesfotmdinA,scleropoides.Thespecimens,someofwhichare
sterileandproducenomatureseed,appeartorepresenthybridsorderiva-
tivesofhybridization,perhapsbetweenA.crassipesandA.scleropoides.These
collectionsareallfromlooseredsandstosandyloamsinsouthernTexasin
Duval,Webb,Hidalgo,andAtascosacountiesofsouthernTexas,withone
collectionfromadjacentTamaulipas.Theyareofsufficientdistinctionto
berecognizedasaseparatenothospecies.
AmarUa.nS.tAh.uTsEXAxS.teWxeebnbsiCso.:Heinnrreidcskasnodna,trnoaodtshiodespo.nHnwoyv..8(3F,ig13s.mi4ND,W5o)f.WTeybpbi,:
16Jul1957,D.5,Correll&l.M.Juhusioii18101(noLOTYPc:LL!;isorvpt:Gil!).
AAmaninthoscleropoick'caulibuserecto-asccndentibusstylis(2—)3etfructibusdehiscentibtis
semiiisseddiffertba.sibusstylaribu.scoliumdistinctum0.2—0.4mmlongumformantibus
(utA.cni.ssipe)etsepali.sflorumpistillatorumvaklereHexisadapices.
Annualherbs10—35cmtall;stemserect-ascendingtodectmibent,with
mm
scatteredobscurelygland-tippedhairsto0.3 long,glabrate.Leafblades
linear-oblanceolate,(5—)13-25(—40)mmlong,2-6(-8)mmwide,obtuse-
emarginatewiththemidveinexcurrentat the tip,narrowlycuneate, the
marginsextendingwelldownthe(3—)5—15(—20)mmlongpetiole,themargins
whitish,crisped,thebladesgreentogray-greenwiththeveinsimpressed
above,stramineousandraisedbeneath.Flowersbornemshort,thickened,
axillarydichasialcymes prodticedateachnodeoftheplantfrom baseto
tip,thecymes3.5—6mmlong,thecymeaxessinuate,stramineous,spongy-
thickened,indurate,0.5—1mmthick,bracteate,thebracts1—1.5mmlong,
green mediallyin adiamondpattern with broad clasping scariousbases,
thestaminateflowersearlydecidtious, thedichasialcymes bifurcatingat
thebasesofeachpistillateBowerandterminatinginpistillateflowers;staminate
flowers: sepals 5, narrowlyovate, 1.5—1.8 mm long,membranousexcept
alongtheexcurrentmidvein;stamens3;maturepistillateflowers:sepals5,
792 SiDA1S(3)
I'Hi. i.l-'li()t()gra|ihsolmaiiirfIrLutandtaU'ccs.A—(!.Aiiiiirinithinsck'r(il>iucles.A—B.I'riiit
wirhinllarL'clstyle-bases,2or^srylc-s((Cameron(x).,Tex.,Ri/ii)o)i2.)97,I')-^- I'l'Liitwith
iion-inllatetlstylebase,notereCLirvedsepals(Mel.eiiiionCo.,'lex.,)'ork4(^)24^),1').l^.A.X
lexensis.MattireIniit;notenarrowstylebase,shar|il\'recurvedsepals(WebbCo.,Tex.,('.orrcll
&joh)isli)>iIHlOl,LL-type).E.A.polygitiioidi^s.Mattirefriiit-se|ials,notesepalshave3veins
andareintlatedandinikiratedatbase(SantoDomingo,lik-iu.n/I}19..KUS).KA.hnnnli\nc/.
MatLii'eealyxwithreflexed,bioadsejials,inlTttedlopoljietKarp(Sonora,Mexico,Bi\///cki^c'e
,v.//.-Tm'i;,I)(;).Cj.a.Uniiii/il/JKiisii.Notespathtdate,erectsepals,intlatedj^encarptop((Cameron
CV>a()n.,DRe//viei)ii/ijiikr28nH(-)H.7(!)'),.AHR.IAZ.).XWihi/iiStn)eic)ibia/r\.iMnatAti=reIcamlmyxahonlddsjielroircaarlilTrtiio;ps.(PimaC^o.,Ariz.,
