Table Of Content.
LEMMA MICROMORPHOLOGY IN THE
ERAGROSTIDEAE (POACEAE)
JESUSVALDES-REYNA
DepartaiNentodeBotanica
VniversidadAutonomaAgrariaAntonioNarro. Buenavista, Saltillo
Coahmla, 23M3. MEXICO
STEPHAN L. HATCH
^.Ai. Tn/cy Herbarium
DepartmentofRangelandEcologyandManagement
lexasA&M Unwersity, CollegeStation, TX 77843, U.S.A.
ABSTRACT
Scanningelectronmicroscopywasusedtoexaminerhelemmamicromorphologyof30
genera and 57 species in the tribe pTagrostideae. Results show four silica deposition
patterns: I)corkcellassociatedwithsilicacell; 2)corkcellsoiitaty; 3)corkcell with
papillae;1)corkcellnotobserved.Thepresenceorabsenceofepidermalpapillae,prickle
hairs,micro-andmacro-hairsisreported.Anelecttonbeamx-raymicroanalysisindicateda
highsilicaconcentrationinallstructuresexaminedincludingthecorkcells.Thedisrribu-
tionofcorkcellsandsilicabodieshastaxonomicsignificanceandtwosubtribeswithinthe
tribeareproposed.
Keyword:lemmamicromorphology,silicadepositionpatterns,taxonomicsignificance,
Eragrostideae, Poaceae
Ki:SlIMHN
Seexamine') lamicromorfologiade la lemade M)generos y 57 especiesde latribu
Eragrostideaemediantemicroscopiaelecrromcadebarrido.Losresultadosmuestrancuatro
patronesdedeposiciondesilice: 1)celulastiberiferaasociadaconcelulasilicifera;2)cclula
suberiferasolitaria; 3)celulasuberiferaconpapila;4)noseobservecelulasuberi'fera. Sc
reporralapresenciaoausenciadepapilasepidermicas,aguijones,micropelosymacropelos.
ElmicroanalisisconhazelectronicodeRayosXindicounaaltaconcentracidndesihceen
todas lasestructurasexaminadas, incluyendolas celulassuben'feras. Ladistribucionde
celulassuberiferasycuerposdesiliceticnesignificadotaxonomicoyseproponendosdentro
delatribu.
Palabrasclave:micromorfologiadelalema,pattonesdedeposiciondesilice,significado
taxonomico, Eragrostideae, Poaceae.
IN'fKOnilCTION
TheEragrostideaetribeiscomposedofwarmseasongrasseswithacenter
ofdistributionin Africa, withextensionstotheIndiansubcontinentand
Australia,andasizeableincursionintoNorthAmerica(Phillips 1982). In
SiDA 14(-1);531-549. 1991.
.
532
NorthAmericathistriLx'isbestrepresentedinthescmiaridsouthwestern
UnitedStatesand northernMexico, whereitmaycomprisemorethan50
percent of the i^'rass vc\^etation ((}ould and Shaw 1983).
hi the United States and Mexico the tribe is represented by approxi-
mately 26 generaand 250 species ofnativeand introduces grasses. The
largestgeneraareI:r^/i^ros!/s. Ai///)/c'Hher<^/a, andSporobol/is. Whereas, there
aretwoorthreegeneraolmediumsize,andtherestofthetribeiscomposed
o[an unusually large proportion ofsmall, often monotypic, genera.
MembersoftheEragrostideaecontainpaniculateinHorescencesthatarc
composedofseveralracemoseorspicatebranches,occasionallyreducedtoa
simplespike. Spikeletscommonlyhave I toseveralfloretsandthereduced
floretswhen presentareusuallyabovetheperfectones. Disarticulation is
above the glumes except in Lya/rns and a few species ofMiihlenbergia
Lemmasare ^-nerved,exceptinSpornholiisandCciLiniovilfa,whichhave I-
nervedlemmas,andinVaseyochliki,withseveral-nervedlemmas,(^aryopses
havealargeembryowithapunctiformorellipsoidhilum, sometimesen-
closed within a tree pericarp(Phillips 1982; GouldandShaw 1983).
Renvoize(1983)surveyed theleafbladeanatomyofthetribeandcon-
cludedthatitsgenerahaveadaptedtopioneerorharshhabitats. Inadapt-
ingtosuchextremestheleafblademorphologyandanatomyhavebecome
highly modified.
Micromorphological features o(the Moral bracts ofgrasses have been
utilized recently as valuable characters that reHects systematic relation-
ships and evolutionary trends. Studies ofthe lemma micromorphology
havebeenreportedbyBjorkman(I960), Hsu(1963),Baum(197I),Clark
and Gould (1973), Thomasson (1978a, 1978b, 1980, 1981, 1984, and
1986),ShawandSmcins(1979),Terrelletal. (1983), WebsterandHatch
(198^),Thompson(1983),andBarkworth(1983).Specificstudiesofsilica
cellandsilicabodieswerereportedbyTerrellandWergin(198I). Inaddi-
tionsilicacellsandsilicabodieshavebeenrecognizedasstructuresoftax-
onomicsignificanceinthegrassfamilybynumerousinvestigatorsinclud-
ingMetcalfe(1960), Ellis(1979), PalmerandTucker(1981),andinother
monocotyledons (Stant 1973).
However, few investigations of the lemma micromorphology of the
wErhaogreoxsatmidienaeedhtahveeebpeiednermmaidseofwigtlhumtehse,elxceempmtaisonaonfdSpaanlcehaeszoi(B1l9e8p3h,rt1d9a84c)h,m
andiWiturihi, and Peterson(1989), Petersonctal. (1989)whoreportedon
the lemma micromorphology and leafanatomy for t2 species ofannual
Mz/hloihcrii^ici. Therefore, ourobjectivewastostudyepidermal featuresof
the lemmaof 30genera, 37 species, and twovarietiesofthistribeusing
scanning electron microscopy. Our specific objective was to document
533
lemma micromorphology and detect different distributions of any
epidermal patterns restricted to specific genera. This intormation would
enable furtherevaluation ofthephylogeny olthe tribe.
mati;riai.s and Mirnions
Lemmasof57speciesrepresentmg 30generaoftheEragrostideaetribe
wereexammed(Table 1)fromherbariumspecnnens(ENCB,TAES,TEX).
Specimenswereselected toberepresentativeoftheirrespectivegenerain
theEragrostideae.AlthoughprimarilyNewWorldgenerawereexamined,
a few Old World genera were included. Three or more specimens per
specieswereselectedfromdifferentlocalities. Allmaterialswereidentified
using the most current treatments ofthe tribe available (Gould 1979;
GouldandShaw 1983). Lemmaswereremovedfrom thefirstandsecond
floretsofmaturespikclets,orientedwiththeapexattheright,mountedon
aluminum stubswith Avery's spot-o-glue to observethe abaxial surhice,
and then coated with 20 nm ofgold palladium inavacuum evaporator.
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534
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,V.uey_lcclu\Na—sh U.S.A. Missohki.BarronCo.:Rig,i^i/i.\723(TAES).
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535
Sampleswereexaminedat 5- 15 kVwith0°tiltonaJEOLJSM-25 SIl
scanningelectronmicroscope. Lemmasofselectedgenerawereexamined
withelectronbeamx-raymicroanalysisontheJEOLJSM-35scanningel-
ectron microscope to determine the elemental content ofspecific struc-
tures.
To examine the effects of herbarium preservation techniques on
specimens, lemmas from living plants {Tndens) were fixed in 2.5%
glutaraldehydebufferedin0.IMsodiumcacodylateforonehour;washed
threetimesin0.IMsodiumcacodylatebufferfor10min.;postfixedin 1%
osmiumtetroxideforonehour;washedthreetimesfor 10min. eachtime
with 0.IM sodium cacodylate buffer; dehydrated in a graded series of
ethanol;driedinaDCP-1criticalpointdryingapparatus;andcoatedwith
TVtubekoatand20nmofgoldpalladium.Lemmasofseveralgenerawere
cleaned inxyleneinanultrasoniccleanerforonehourtoremoveepicuti-
CLilarwax.
Rl-SUL'l'S AND DISCUSSION
Themicromorphological surfacefeaturesofthelemmaoftheEragros-
tidcae exhibit typical "chloridoid" characteristics (Prat et Vignal 1968),
suchaspapillae,microhairs,macrohairs,abundantpricklehairs,andsilimca
cells.ThiscorrespondswiththeconclusionreportedbyRenvoize(1983)
theanatomicalsurveyoftheleafbladeofthistribe. Uniquesilicadeposi-
tion was observed in cork cells (Figs. 1-2). An electron beam x-ray
microanalysisofthisstructureindicatedahighconcentrationofsilica.The
analysisofthecorkcellforErioneimmaimaceum(Fig. 3)showsthatahigh
silicaconcentrationisassociatedwithsomeartifactsofsamplepreparation
(gold&palladium), andelectronsemittedfromthesurroundingenviron-
mentwithin the microscope(copper).
For comparative purposes, since all the samples were obtained from
driedherbariumspecimens, freshlemmamaterialfromTridensstrictuswas
analyzed(Fig. 4).Theosmiumconcentrationwasremarkablyhighdueto
theosmiumtetroxideusedinspecimenfixation.Theanalysesoftheprickle
hairs and thesilicacell (Figs. 5-6) from thesame lirwneuronavenaceum
specimenusedforthecorkcellanalysis,showsahighsilicaconcentration.
The presence or absence ofthe cork cell, silica cell, papillae, prickle
hairs,microhairs,andmacrohairsindicatefourdistinctivepatternswithin
the tribe. Thefourpatternsarediscussed with representativeexamples.
I. Corkceil associated with silicacell. In Figure 7 thecorkcell is
evident with the associated round silica cell in Trioclia irritam var. lax-
ibseplilca-tsah.apIendTsrilwidciaaceplmlsgeannsdathnedaNseseorcaigartoesdticsorrekptcaenlsl,arreelsathiovwelny(sFhiogrst.d8u-m9b)-.
Kidney-shapedsilicacellsandassociatedcorkcellareevidentinEragrostis
536
erosci and E. muxicana (Figs. 10- 11). The cork cell, dumbbell-shaped
silicacell,andpricklehairsof£.cHums(Fig. 12)exhibitasimilarpattern
reported by Baum (1971) m lemmas ofArm/, and Terrell and Wergm
(1981) in Zizama.
II. Cork cells solitary. This pattern characterized a number ofthe
generaobserved. Infivespeciesoi'Tmknstheshapeofthecorkcellvaries
from crescent or kidney- to fattened dumbbell-shaped (Figs. 13-17).
Trtphisis(iMg. 18),Sohmui(Fig. 19),Ele!^swe(Fig. 20),andDactyloctenum
(Fig.21)havesilicacellsinrowsthatarenotassociatedwiththecorkcells.
III. Corkcellspapillate. Inthesetaxathecorkcellisassociatedwith
smallroundedpapillae,similartotheonesillustratedbyClarkandGould
(1975),Thomasson(1978b),andTerrellandWergin(1981). Thispattern
is seen in l.eptocbloa(Fig. 22), Tnpogini(Fig. 23), Leptoatrydinn(Fig. 24),
Gymnopoii^ou (Fig. 25), Tnchonaira (Fig. 26), Vaseyochloa {V\^. 27), and
Sderopo^on(Fig. 28). Whenasilicacellwasobserveditwasassociatedwith
thecorkcellandpapillae. InGouniuiacorkcellandpapillaearcshownbut
prickle hairs are not abundant (Fig. 29). Erioneuron seems to be in-
atsesromceidaitaetdewbitehtwteheensiplaitcatecrenll1(aFingds.\\3,1si-nc3e4)t.heThcoerskhcaeplelomfatyheorcomrakycenllotwabse
characteristicallyelongatedvertically inllnmh'/iroii, withanoutlinevary-
ingbetweenoblongtocrenateorscalloped. Anabundanceofpricklehairs
occurs along theepidermal lemmatae.
35I-V.3C6):.orTkhceellmsicnroothaoibrsserobvseedr.veTdhiwserpeathtecmrimsspwhaesrifcouannddisnimMi/lnairrotao(tFhiogss.e
reported by Sanchez (1984). Similarmicrohairs were also present in the
genusEriomiiron. Papillaewerealsoevidentandabundant. Bkphandachm
a3n7d-Re3d8j)i.ddAibctunhdavaentabpurnidckalnethpariircsklheavheaibresetnhrroepuogrhtoeudtftohreHehppihdaerrimdiasch(nFeigbs.y
Sanchez(1983). InPereilenni thepricklehairsarcassociatedwithpapillae
overlongcells(Figs. 39-4()). However,acleardistinctioncouldbemade
based on the long cell's shape. Peralef/uiciliutum hascharacteristic raised
cell walls forming a ridge around the long cells. In Trnaochloapapillae,
dpraincckeleos(asnmdalplithtoeodklsonargecselelesnarinesSphoorowlnml(uFsigo.zcAtr1)k.cMwmac(rFoihg.ai4r2s).anBdleapnhaarbounne--
iirouirnholcpisandMNhlenhtr^i^iamierskyihavebiccllularmicrohairs,prickle
hairs, papillae,anddeeplyundulatinglong-cellsmargins(Figs. 43-44).
Thischaracteristicshapeoflongcells isalsoobserved inChahoissaea(Fig.
45),Lyi//r//.\pbleoidts(Fig. 46),M/djlenha-i!^iciuiniKtnstnui(Fig.47),andM.
jctitiii^hitci (Fig. 48). Dumbbell-shaped silica cells with a relatively wide
centralportionareseenin^\.plNmbm(Fig.49),SptmMNspyuDuidatHS(Fig.
50), S. airoides(Fig. 51),S. asper(Fig. 52), .V. ayptandriis(Fig. 53), and
Crypsis(Fig. 54).
337
^mi\,-^5i,'»^|*'
n
FIC.S.1-6.Lemmamitromorpholot^yaiulijniphsofclcctronbeamx-r.iyinKroanalysisoflinimeimm
andlYidois.Norethatahii^lisilitaconcentratuMiwasiouiiil(orallstructuresanalyzed.Fij;. I.Silica
bbmpPorieddicacirmkeo=lsaenx/-aph;rl.aaaiylryisl.iiafmcsdFiiinloctli;trn.mioc;a6Con.rnpaknShltiyclosei=klic^lasaopc.o{/erf.li«Il.ctm)koh./lfeecF£icm.sho;ar.<Mikr»r2;nw.ctwesulc(csldtomio=sl«Veat/(/si;«i.Apal.Yiai-c/irRafte.rsucieM6l,lJti,l9c'«,-au.?)wcc)ce.Fliil.lA=ui.aumn/Vcad=o-raF6ksgl2seo9occle2ctldi)rla.;;otCneSFudIib,e^c=.=oarm4ck.soxipcl-Eepirllcelaear.y.c;tFmrUiiogscn.ro=b.a^.enoaaFlsmlymescixit-usrrmoao;nyf
538
MCrS.7-12.SEMpli()t()micr()^urapli.s()tlemmasurfacesfromsclecr«l^C'iu-raofHm.yrosriclcac.Note
rhatthesilicacellsvaryfromshortdumbbell-tokidney-shapedandareassociatetlwirhacorkcell.Fi^.
7,'irimluiiryitu)!^var,hixnpuala{lit:ndm,m\\\).Fi^.H.'l.piin^ms(Hubbardl]')S).Fi^».9.Heenigroitts
r\L2--Chn/i;'s.{tCi,/ni:„/LnIi1(1Si2h4vi1ti).-l-4i0i;6.5)1.0.SECru,v,n,>sii!l>ic,armcue1(1«;),c-cm\=6)c.orFkigc.ell1;1j.iCh,=mexpuraiichklle(Hhaaticr.he!al.-f5H8).Fig.
539
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510
I'lCiS. I9~2I. LemmaepKlcrm;ilpartcrnstorstkcadSfucniolLragrosridoie. Notediesmall
plaUpiIl)laewiarshsoseiiahtceadcwelulls,miheroewosrka.nedllnaonrdarshseocsilaetnedderw,ivtdhloauscomrakerceolhla.ir|s•,.,^.Fi2.0y..1E9l.cSmaihmmut,ml[uiluiH(u/,im(vS/o/h«mt
c'/e'^l'«^l-.;SJ-ct)^.j-l''v\ctM;o".r"/k2-V1t'.e/l<Dl'M;,t7/p^/al^m.t=(a/inpi<a/up/wicl,.l\ialve-^;JyJp/mi,iaSu)m,={IS-,dt,ymn.aatd2r)ao.-haU/i.rM<.9/»'t)o/.n'.F/./g>/.;/11r/.///U7pw.t,/orc/,h/l;/uu(Hi./i/mrm/^rru,(Lmuscnd=21si]liIc)a
