Bol. Soc.Argent.Bot.38(1-2): 5-12. 2003
ULTRASTRUCTURAL STUDY
OFSTROMBOMONAS
(EUGLENOPHYTA)FROMTHE
PROVINCE
OFCHACO
(ARGENTINA)1VISITACION CONFORTI2 andYOLANDAZALOCAR DE DOMITROVIC3
Summary:Nine taxa of StrombomonasDeflandre (Euglenophyta) fromthe province of Chaco (Argen¬
tina) have beenexaminedwithscanningelectronmicroscopy (SEM). Details ofthe lorica ultrastructure
are described and illustrated, five ofthem for the firsttimewithSEM:Strombomonas confortaZalocar de
Domitrovic, S.eurystoma(Stein)Popova, S.longicauda(Swirenko) Deflandre, S.pizarroi Zalocar de
DomitrovicandS.urceotata(Stokes)Deflandre.For the other taxa (S.tellii Zalocar de Domitrovic, S.
scabravar.hiperintermedia Conforti, S.dípteraZalocar & Tell and S.tetraptera Balech&Dastugue) additionalcommentson relevant morphologicalcharacteristicsare furnished.
Key words: taxonomy, ultrastructure,Euglenophyta, Strombomonas,Chaco,Argentina.
Resumen:Estudioultraestructural de Strombomonas (Euglenophyta) de laprovinciade Chaco(Argen¬
tina).Enestetrabajose examinaron con microscopio electrónicodebarrido (MEB)nueve taxones del
genero Strombomonas Deflandre (Euglenophyta),colectadosen cuerpos deagua dela provincia de| Chaco(Argentina).Se describeeilustrala ultraestructurade suslorigas,de lascualescincoson analizadaspor primera,vez con esta técnica: StrombomonasconfortaZalocar de Domitrovic, S.eurystoma
(Stein)Popova, S.longicauda(Swirenko)Deflandre, S. pizarroiZalocar de Domitrovic y S. urceolata (Stokes) Deflandre.Para lostaxa restantes(S. telliiZalocarde Domitrovic, S. scabravar. hiperintermedia Conforti, S. dípteraZalocar & Telly S. tetraptera Balech & Dastugue),se amplía sudescripciónyse detacan algunas diferenciascon observaciones previas.
Palabrasclave: taxonomía, ultraestructura, Euglenophyta, Strombomonas, Chaco, Argentina.
INTRODUCTION
Trachelomonaswereelevatedtothe generic levelby Deflandre (1930)and renamed Strombomonas. Thisauthorbased the separation of thetwo gen¬ era onratherpoorlyand doubtfully definedmor¬ phological characters ofthe envelope. Conse¬ quentlysomeresearches have discussed the adop¬ tion of Deflandre’s classification (Balech, 1944; Dunlapetal., 1986;Tell & Conforti, 1988; Conforti &Nudelman, 1997). This controversial group has been scarcely studied around the world. In Ar¬ gentinatheywerealso rarely found and the high¬
estnumber ofreportscorrespondtoourstudyarea (Tell, 1998; Tell & Zalocar deDomitrovic,1985; Zalocar deDomitrovic,1991). Some ofthese speci¬ mensshowed suchanoriginal and differentmor¬ phology thatthe authors considered themas new taxa. Although in previous papers (Hager,1979;
-Tell & Conforti,1984,1985, 1988;Dunlap eta!., 1986;Conforti, 1993; Confortielal.,1994;Couté
& Thérézien,1994;Conforti & Pérez,2000) some species have been described with scanning elec¬
tronmicroscopy (SEM),there are still very few
taxa studiedat this level. Thepresentinvestiga¬
tionattempts toimprove this situation. The Argentine northeastern region isveryrich
in vegetated waterbodies (specially inthe Paraná River floodplain)whichpresentsfavorablecondi¬ tions for thedevelopment of euglenoids. Numer¬ ous taxahave been reported for this area(Tell, 1980, 1998; Bonettoetal., 1984;Tell & Conforti, 1985;Tell & Zalocar de Domitrovic, 1985; Zalocar de Domitrovic etal., 1986; Zalocar deDomitrovic,
1991, 1993,1999,inpress; Conforti, 1998,1999). Amongthese,ahigh number of species belong¬
ingto the genus Strombomonas Deflandre were described. Initially, severaltaxaof this genus,were classifiedasspecies of Trachelomonas Ehrenberg. Duetothedifferencesinlorica morphology,spe¬ cially for its sac - shape, members of the Trachelomonas subgroup “Saccatae” 'of
’Dedicado
al Prof. Dr. Sebastián A. Guarreraenocasión desu 90° aniversario.2Dpto.
deBiología,Facultad-deCienciasExactasy Natu¬rales, CiudadUniversitaria,Pab. -II,1428,Buenos Aires,
Argentina,[email protected]
3Centro
deEcologíaAplicada del Litoral -(CONICEf),C.C.291,3400, Corrientes, Argentina,[email protected]
Lorica 50-60 pm long,23-26 pmwide, rhomboi-dal.Middleregioncircular inapicalview.Anterior Thematerials analyzedinthis workwereobtained end narrowed intoalong cylindricalneck (10-14 pm
fromdifferentwaterbodies of theprovince of Chaco, long),widened,obliqueandirregulartowards thetip northeasternregionofArgentina (27° 27’ S; 58° 49’ (6‘7 P™ wide). Posterior endabmptly narroweddown W),whichwereshallow and vegetated withmacro- t0aconical,straight,verylongcauda,20-22 pm. Wall phytes. Theywerecollectedfromthefollowingplaces: yellowish,thick,rough,withnumerous adheredpar-E1 PuenteLake,CuatroDiablosLake,“A” and “B” ticlesonits surface. Europe (Huber-Pestalozzi,1955). LakesinPresidencia Roque Sáenz Peña, Cochereck InSouthAmerica: Argentina,Buenos Aires (Conforti,
Lake and Felman Lake (seemapofthe studyareain 1981),ilis recorded in Chacoforthe first time.
Zalocar deDomitrovic,1991). Water was
sieved
MATERIAL
ANDMETHODS
Material: ARGENTINA: Chaco,Lake Felman, 5 througha25pmmeshplanktonnetand fixed with4% Nov. 1978,Zalocar deDomitrovic,C-855.
formaldehyde.For SEMobservations,organismswere
isolated underadissecting microscopewith the aid of Strombomonas pizarroi Zalocar deDomitrovic,
micropipettes,dehydratedinaseriesof ethanolsolu¬
tions(50to100 %),air driedonaluminum foil and pjgS, 2,A-C coated with gold / palladium. Specimenswere exam¬
ined andphotographedwithaPhillips 505 SEMatthe
ElectronMicroscopyServiceofCITEFA.Sampleswere p0resurroundedbyashort neck (3-3.5 pmlong), wid-deposited inthe PhycologycalCollection ofthe ened,oblique and irregularatthe distal end (5-6 pm Centrode EcologíaAplicada delLitoral(CONICET), wide). Posterior end abruptly narrowed downtoa
conical,straightoroblique,longcauda(18-20pmlong). Wallthick,coarse,withvery irregular contour,totally
covered withagglutinated.exogenousmaterials. Among theseparticles,itispossible todistinguish somewithlargedimensions. This observationis op-StrombomonasconfortiiZalocar deDomitrovic, positetothe originaldescriptionofthisspecies, where Cryptogamie,Algol.12 (4):273-274,fig.1a-d, the author said that the loricasurfaceissmooth. This
specieswashas only been recordedinmaterialsfrom Chaco,Argentina (Zalocar deDomitrovic,1991).
Cryptogamie, Algol. 12(4):274-276,fig.2a-b,1991.
Lorica 48-55pmlong,28-31pmwide,pear
-
shaped.Corrientes.
TAXONOMICAL
DESCRIPTIONS
1991. Figs. 1, A-D
Materials: ARGENTINA: Chaco, Lakes “A”, 23 Dec.1989and Lake “B”inPresidenteR. Saenz Peña, 24 Set. 1990,Zalocar deDomitrovic,C-3507,C-3561
.
Lorica 36-38pmlong,28-30pmwide,bell
-
shaped. Anterior end narrowed intoacylindrical neck (4-4.5x 6.5-7 pm),obliqueorirregularatthetip.Posterior end flattened surrounded byawing-
shapedexpansion (3-4pmlong),irregularinthe free edge (Fig. 1 A-B,arrow¬ heads).The caudais conical,straightorslightlyob¬ lique (13-15x7-8.5 pm). Wall totallycoveredwithag¬ glutinated exogenous particles. The dimensions oftheloricaexamined in thisstudyweresmallerthan those
analyzedinthe originaldescription, 47-48x38-38.5pm. Thisspecies has only been foundinmaterials from
Chaco,Argentina(Zalocar deDomitrovic,1991).
Strombomonas tellii Zalocar de Domitrovic,
Cryptogamie, Algol. 12 (4):276,fig. 3 a-c, 1991
.
Fig. 2,DLorica 49-51pmlong,31-33 pmwide,rhomboidal,
withanequatorial fold;broadly elliptic inapical view. Anterior end prolonged intoashortcollar(3-4 pm long),widened, obliqueorirregularatthe distalend
(7-8.5pmwide).Posteriorendnarrowedintoastraight Material: ARGENTINA:Chaco,Lake “A”in conical cauda (10-11 pmlong), straight and symmetri-Presidente R. S. Peña, 23 Dec.1989, Zalocar de caloroblique and asymmetricalinlateral view. Wall verythick,coarse, yellowishtolightbrown, shows exogenousparticles coating its surface. Argentina
Strombomonas longicauda (Swirenko) Deflandre, (Chaco,Zalocarde Domitrovic, 1991) and Brazil Arch. Protkde. 60 (3): 599-560, figs.108-109,1930. (Conforti, 1993).
Figs.1,E-F
Domitrovic,C-3507.
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Fig. 1. A-D: Strombomonasconforta,A, C: generalviews;B:detail of theposteriorendshowingadetail of the wing- shaped
expansion (arrowhead); D: detail of the neck and loricasurface;E-F: Strombomonaslongicauda,E. detail of the collar apd loricasurface;F:generalview. Scale bar valuesaregiven inpm directlyonthemicrographs.
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Fig. 2. A-C: Strombomonas pizarroi, A. general view; B. detail of the neck and loricasurface;C. detail of theposteriorend; D:Strombomonas tellii,generalview;E. Strombomonaseurystoma,generalview;F:Strombomonas urceolata, generalview. Scalebar valuesaregiven in pm directlyonthemicrographs.
Presidente R.S.Peña., 23 Dec. 1989,Zalocar de tides. Thistaxonhas been already described with SEM byTell & Conforti (1988), but theyonlyhave reported specimens like those shown in Fig.3 B. Strombomonas eurystoma(Stein) Popova,Flora- Strombomonasscabravar.hiperintermediawasonly sporovychrastenij SSSR, 8, pg.:206,PI.XX, recordedin materials fromArgentina,BuenosAires (Conforti, 1989)
and
Chaco (ZalocardeDomitrovic,1991).
Domitrovic,C-3507.
figs. 1-11,13. 1966.
Fig.2,E
Materials: ARGENTINA: Chaco, Lakes
Cochereck,9 May 1979,and “B” in Presidente R. S. Peña, 24 Set. 1990, Zalocar de Domitrovio, C-1092, C-3561.
Lorica 43-47.5pmlong,20-24pmwide,irregu¬
larlyellipsoid,compressedinlateral view. Pore sur¬ roundedbyashort neck (2-2.5x5-5.5 pm),undulated
atthefreeend.Posterior endnarrowed intoaconical
cauda(7.5
-
9.5 pm long), roundedatthe tip. Wall yellowish,thick, totallycoveredwith agglutinated exogenousmaterials.Widespread(Huber-
Pestalozzi,1955).InSouthAmerica:Argentina,Corrientes (Tell, 1980),recordedin Chaco for the
first
time andBrazil (Conforti, 1993).Strombomonasdíptera
Zalocar
&Tell,Bull. Mus. Hist. Nat., Paris 1:42, lam.VIII,fig. 10. 1980.Figs.3,E-F
This species is characterized bythepresenceof twohelycoidalhelicoidal wings 2-3 pmlong(Figs. 3
E-Material: ARGENTINA: Chaco,LakeElPuente, F, arrowhead), that surround thebody fromthe collar to
the
cauda. Thestudiedspecimens presentedsome differencesthanwith those described with SEMbyStrombomonas urceolata (Stokes) Deflandre,Arch. Couté &Thérézien(1994).The dimensionsaresmaller;
5 Dec. 1985,Zalocar deDomitrovic,C-312.
lorica30.5-42pm long, 19.5-21pmwide,neck 4-7pm long,3.5-4 pmdiam.,cauda 3-4pm.Theenvelope showed very irregularcontour.Itis extremely thick, Lorica 38-55 pmlong,20-26 pmwide,urceolate, coarse, withnumerousexogenousparticles adhered
slightlyflattenedinlateralview. Sidesstraight,con- onit its surface which increasethe thickness ofthe
vergingtothe anterior endlikeacylindricalshortcol- wingsthickness. This specieswasonlyrecorded in
lar (4-4.5x5-5.5pm),widened,obliqueorstraightat materialsfromSouthAmerica;Argentina,
Corrientes
thedistal
end. Posterior endabruptly taperingtoa (Tell, 1980),Chaco(Tell& ZalocardeDomitrovic,1985) conicalcauda(8-9 pm long). Walltotallycoveredwith andBolivia (Couté &Thérézien,1994).exogenousagglutinatedparticles.
Widespread
(Huber-Pestalozzi, 1955).InArgentina,Corrientes (Tell, 1980) andChaco (Tell &Zalocar deDomitrovic,1985).Protkde.60(3):586-587,figs. 70, 71, 73.1930. Fig.2, F
Material:
ARGENTINA:
Chaco, Lake Felman, 5 Nov.1978,Zalocar deDomitrovic,C-855.Material: ARGENTINA: Chaco, Lake Cuatro Strombomonas tetrapteraBalech & Dastugue,
Diablos,24April1980,Zalocar deDomitrovic,C-333. Physis12: 355. 1938.
Figs.3, G-H Strombomonasscabra var.hiperintermedia Conforti,
Cryptogamie, Algol. 10(1): 74-75, figs. 14a-c. 1989.
Figs.3, A-D
Lorica fusiform, quadrangular withconcavesides
inapical view. This species ischaracterized bythe presence of fourhelicoidal wingsthatsurround the Lorica 55-65 pmlong,19.5-33pmwide,ellipsoid, body fromthecollartotheposteriorend.Neckshort,
Neck cylindrical,long(10-13 pm), denticulateatthe wide andirregularatthetip.Posterior end gradually distal end and surroundedattheproximal end bya narrowedintoaconical short cauda (Fig. 3H). The welldevelopedannular externalthickening(2-3.5x9- wall is totally covered by agglutinated exogenous 13 pm),whichshowsirregularcontour(Fig. 3 C,ar- granules,whichproduceaveryirregularenvelope
rowhead).Posteriorend graduallytaperingtoalong surface.Thespecimensstudied in this workshowed conicalcauda,thatiseither long (11-13pm,Fig.3 A) smallerdimensionsthan thosedescribedbyTell & orshort (5-6pm,Figs.3B,D). Wallbrowntoreddish, Conforti(1988).
These
authorsreportedlorica 43-66Á
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Fig. 3. A-D: Strombomonasscabravar.hiperintermedia,A-B. generalviews;C.detailofthe neckshowingannular thickening (arrowhead) and lorica surface; D. detail of theposteriorend; E-F: Strombomonasdíptera,generalviews showing the lateral wing (arrowheads); G-H: Strombomonas tetraptera,G.generalview,H. detail of theposteriorendshowingthe helicoidalwing (arrowhead) and lorica surface. Scale bar valuesaregiven inpmdirectlyonthemicrographs.
whereasweobserved lorica 44-47x18.5-26um,col¬ lar 3-3.5x5-5.5pm,withalongercauda,7.5-8x6-6.5 pmand wings with different thickness,4-5 umin the firstgroupand 1.5-2pmin the latter. In spite of the observed morphological differences,wedonotesti¬
matethat there is enough evidence enoughtocon¬ sider these organismsas two separate taxa.Size varia¬ tionsareprobablyproduced bytodifferentamounts of agglutinated exogenous material. On the other
hand,the observed organisms do notcoincide ei¬ ther with those described as Strombomonas tetraptera var.gallica (Bourrelly & Couté, 1978) which donot presentwings. This species has exclu¬
sively been found in South America: Argentina, BuenosAires(Balech&Dastugue, 1938),Chaco(Tell &Zalocar de Domitrovic,1985),Brazil(Conforti, 1993) and Uruguay (Conforti & Pérez,2000).
Materials: ARGENTINA: Chaco, LakeFelman,5 Nov.1978 and“B”in Presidente R.S.Peña, 24 Set. 1990,Zalocar de Domitrovic, C-855, C- 3561
.
(Uherkovich & Schmidt 1974; Yacubson 1980; Tell & Zalocar de Domitrovic 1985; Thérézien 1989; Zalocar de Domitrovic 1991; Conforti 1993; Couté & Thérézien 1994;Menezes 1996;Tell1998)who studied
eugle-noids from different places in SouthAmerica,have described several species of Strombomonas exclu¬ sively found in this continent. Thesereportscould indicate thepresenceofendemictaxaorprobably they could be the result of the few studiesonthisgenusin otherplaces of the world. Further analysis of samples fromother localities should enhanceourunderstand¬
ingnotonly ontheir distribution butultimatelyon thedifferentcharacteristics of thispoorlyknown group of euglenoids.
ACKNOWLEDGEMENTS
This workwassupportedby thegrantsPICT6017/ 99and UBACYT X 092. The authorsareindebtedto
Mr. D.Gimenez forhis assistance with the SEM (CITEFA).
CONCLUSIONS
BIBLIOGRAPHY
Amongthe studiedtaxa, S.eurystoma and S.longicauda are newrecords for the province of Chaco. S.conforta,S.eurystoma,S.longicauda, S. pizarroi and S. urceolata aredescribedwith SEM forthe first time.In spite of the fact thatS.díptera, S. scabravar.hiperintermedia and S. tetrapterahave been already described with SEM in previous works,
weincludeour observations becausethey showed morphological differences with those materials.
Ontheotherhand,ourSEMobservationscon¬ firm that the characteristic loricae coarse, irregular andgranulate in thisgroup arecaused by thepres¬ enceof agglutinated exogenous particlesonits wall. All thestudiedtaxahave shown different materials adheredonits their surface. Tell & Conforti(1988) reported thatthis condition could be usedasagood valid taxonomic charactertoseparateStrombomonas Deflandre and Trachelomonas Ehrenberg, but pos¬ teriorly Conforti & Nudelman (1997) reportedone species of this latter genus with adhered material.
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Recibido el 19 deFebrerode 2003, aceptado el 22 de Abril de 2003.