Bulletin of the Botanical Society of Argentina

Bulletin of the Botanical Society of ArgentinaOn-line versionISSN1851-2372

Bull Soc Argent.Bot.Crdoba v.45 n.1-2 Jan. / June.2010

Paleobotany And PALYNOLOGYPalynological studies of Argentinean species of the generaCereus,Cleistocactus,Denmoza,EchinopsisandMonvillea(Cactaceae, Cactoideae)

Elsa LATTAR1,2& Graciela Square1,31Chair of Palynology, Faculty of Natural Sciences and Surveying, Universidad Nacional del Nordeste, Corrientes, Argentina.E-mail:elsalattar@yahoo.com.ar;elsilattar@gmail.com2Northeast Institute of Botany, IBONE3Center for Applied Coastal Ecology, CECOAL

Abstract: The pollen morphology of 19 species belonging to five genera (studiedCereus,Cleistocactus,Denmoza,EchinopsisandMonvillea) of Cactoideae (Cactaceae) in Argentina subfamily were described using light microscopy and scanning electron microscopy:Cereus aethiopsHaw .,C.argentinensisBritton & Rose,C.forbesiiCFFrst Otto ex.,C.haenkeanusFAC Weber ex K. Schum.,C.stenogonusK.Schum.,C.uruguayanusR.Kiesling;Cleistocactus baumannii(Lem.) Lem.C.hyalacanthus(K. Schum.) Gosselin,C.smaragdiflorus(FAC Weber) Britton & Rose;Denmoza rhodacantha(Salm-Dyck) Britton & Rose,Echinopsis ancistrophora. Speg,E.aureaBritton & Rose,E.leucantha(Gillies ex Salm-Dyck) Walp.,E.mamillosaGurke,E.oxygona(Lynk) Zucc.,E.rhodotrichaK.Schum.,E.tubiflora(Pfeiff. & Zucc.)Monvillea cavendischii(Monv.) Britton & Rose andM.spegazzinii(FAC Weber) Britton & Rose.The pollen grains of the species analyzed are estenopalnicos mainly tricolpate, subesferoidales to prolate, with perforated tectum, espinulado I microespinulado or nanoespinulado.Finding small differences in shape and size.Pollen Type: two pollen types were determinedCereus argentinensisand Pollen Exchangemamillosa Echinopsis.A key to identify and distinguish the above types is presented.Keywords:Polen; Morphology; Cactaceae; Taxonomy.Summary:palynological studies of the species in Argentina generatesCereus,Cleistocactus,Denmoza,EchinopsisandMonvillea(Cactaceae, Cactoideae).Pollen grains morphology of nineteen species generates Belonging to five (Cereus,Cleistocactus,Denmoza,EchinopsisandMonvillea) of the subfamily Cactoideae (Cactaceae) in Argentina, are Described using optical and scanning electron microscopy:Cereus aethiopsHaw.C.argentinensisBritton & Rose,C.forbesiiCFFrst Otto ex.,C.haenkeanusFAC Weber ex K. Schum.,C.stenogonusK.Schum.,C.uruguayanusR.Kiesling;Cleistocactus baumannii(Lem.) Lem.C.hyalacanthus(K. Schum.) Gosselin,C.smaragdiflorus(FAC Weber) Britton & Rose;Denmoza rhodacantha(Salm-Dyck) Britton & Rose,Echinopsis ancistrophoraSpeg,.E.aureaBritton & Rose,E.leucantha(Gillies ex Salm-Dyck) Walp.,E.mamillosaGurke,E.oxygona(Lynk) Zucc.,E.rhodotrichaK.Schum.,E.tubiflora(Pfeiff. & Zucc.) andMonvillea cavendischii(Monv.) Britton & Rose,M.spegazzinii(FAC Weber) Britton & Rose.The pollen grains of the species are stenopalynous Analyzed, mainly tricolpate, subspheroidal to prolate, tectum microperforate, equinulate, microechinate or nanoechinate.Little Differences in form and size were found.Two pollen types Could be established: pollen typeCereus argentinensisand pollen typemamillosa Echinopsis.A key to identify identity and Distinguish the pollen types was given.Key words:Pollen; Morphology; Cactaceae; Taxonomy.

INTRODUCTIONCactaceae family is a native of the American continent, represented by 100 to 150 genera and over 2000 species typical of arid regions, but also inhabit rainforests and temperate-cold areas (Kiesling, 2005).In the Catalogue of Vascular Plants of the Southern Cone 41 genera and 412 species for Argentina (cited Kieslinget al., 2008).This group of plants includes herbs, shrubs and succulent trees, mainly distributed in the phytogeographic provinces of Chaco, the Espinal, the Monte Prepuna and in which they are dominant.Many species are cultivated as ornamentals, others as food for its fruit, the ethylene used as wood and some species such as medicinal (Kiesling, 2005).In this study, pollen species belonging studied five genera represented in corresponding to the Argentina Cactoideae subfamily:Cereus,Cleistocactus,Denmoza,EchinopsisandMonvillea.publications referred pollen Cacti are scarce in Argentina.Internationally can cite the work of Kurtz (1963), who analyzed the pollen of some species ofCleistocactus,DenmozaandEchinopsis,genres treated in this work.Nowicke (1975), in his work concerning pollen by Centrospermales examined 12 species of Cactaceae.Leuenberger (1976) described the pollen morphology of Cactaceae and its taxonomic significance, studied more than 600 species belonging to almost every genre of the family of six of them which are treated in this work.During recent years, were carried out palynological studies of different subfamilies and genera Argentines.Among them we can mention: Garralla & Square (2007), who described the pollen morphology of the generaAustrocylindropuntiaBackeb,Maihueniopsis. Speg,OpuntiaMillandTephrocactusLem.(Cactaceae, Opuntoideae), stating that these genres grains are spheroidal, nonpolar, and pantoaperturados radiosimtricos, also found differences in the type of exine, shape and number of pores and the presence of supratectales elements among different taxa.Garrallaet al.(2008) analyzed the pollen morphology of the speciesRebutiaK.Schum., Mainly finding differences in the type of openings.Then Garralla & Salgado (2008) studied pollen speciesRhipsalisGaertn.of Argentina and established differences in the number of openings and the arrangement of the same on the surface of the pollen grains.Finally, a palynological study of the subfamilies was publishedMaihueniodeaeandPereskioideaeof Argentina & Garralla made by Square (2009), who could distinguish the two subfamilies by the number and position of the openings.This work is part of a larger project that involves knowledge pollen almost all species of Cactaceae from Argentina.These results provide data of diagnostic value for the family.It will also help with the 4th vol.Flora of the pollen Northeast Argentina, which is under preparation.MATERIALS AND METHODSThe samples used for the realization of the palynological preparations were obtained from specimens deposited in the herbarium of the Institute of Botany Darwinion (SI), the Institute of Botany of the Northeast (SELV) and Instituto Miguel Lillo (LIL).were studied pollen samples 1-4 specimens of each of the species.ForCleistocactus yalacanthusonly became available a copy of Bolivia, therefore further examined a specimen grown in the Botanical Garden Berlin-Dahlem (Germany) from specimens, wild individuals of Argentina.preparations palynological were performed the acetolysis of Erdtman technique (1966), the material was mounted in sealing them with paraffin glycerinated gelatin.Such preparations are deposited in the Palinoteca of Northeastern University (PAL-CTES).Observations and measurements were performed in 20 pollen grains for each of the species under study.Using an Olympus BX 40 were analyzed: polar axis, equatorial diameter, shape, position and number of openings, area, symmetry, thickness and exine sculpture.The photographs were taken with a digital camera Canon Power Shot S55.The observation and interpretation of the sculpture was completed with the observations and photomicrographs obtained with a scanning electron microscope (SEM) Jeol JSM-580 OLV of the Ministry of Science and Technology UNNE.Previously metallized oropaladio acetolizado and pollen material was used.terminology suggested by Nilson & Praglowski was used (1992), also being consulted Puntet al.(2007), for the name of the spines according to their different heights.Material examinedCereus aethiopsHaw.ARGENTINA.ProvSkip: Dept. General Cafayate,04.I.1972,Krapovickas et al.20621 (SELV), PAL-CTES 7019;ProvCorrientes: Dept. Capital, 18.XII.2003,Meza Torres163 (SELV), PAL-CTES 7403.Cereus argentinensisBritton & Rose.ARGENTINA.ProvCorrientes: Dept. Lavalle,23.XII.1979,Schinini et al.19064 (SI), PAL-CTES 7020.Cereus forbesiiFrst CF Otto ex.ARGENTINA.ProvFormosa: Dept Laish,10.XII.1999,Di Giacomo438 (SELV), PAL-CTES 7407.ProvSalta, Dept. Rivadavia,03.III.1984,Schinini,19583 (SELV), PAL-CTES 7406.Cereus haenkeanusFAC Weber ex Schum.ARGENTINA.ProvJujuy.Dept. San Pedro, San Pedro to Santa Clara, 11.XI.1995,Kiesling et al.8719 (SELV), PAL-CTES 7413.Cereus stenogonusK.Schum.ARGENTINA.ProvCorrientes:Dept.Capital,25.X.2003,Meza Torres. 161 (SELV), PAL-CTES 6808Cereus uruguayanusR.Kiesling.ARGENTINA.ProvCorrientes: Dept. Capital,12.XII.2003,Meza Torres164 (SELV), PAL-CTES 7305.Cleistocactus baumannii(Lem.) Lem.ARGENTINA.ProvEntre Ros: Dept. La Paz,Step Yunke, 03.XI.1998,. Bacigalupo et al1666 (SI), PAL-CTES 7022;ProvFormosa: Dept. Laish,18.XI.1999,Di Giacomo432 (SELV), PAL-CTES 7408.BOLIVIA.Dept.Tarija, 20.XII.1994,Zuloaga et al.1365 (SI), PAL-CTES 7024.Cleistocactus hyalacanthus(K. Schum.) Gosselin.ARGENTINA,ProvJujuy.Schwerdtfeger17649 (B) Det.Leuenberger.This specimen is cultivated from wild specimens.BOLIVIA.Dept.Tarija: Prov Aniceto Arce, undated,Beck et al.22149 (SI), PAL-CTES 7023.Cleistocactus smaragdiflorus(FAC Weber) Britton & Rose.ARGENTINA.ProvCatamarca: Dept. Capital,undatedUlibarri895 (SI), PALCTES 7025.Denmoza rhodacantha(Salm-Dyck) Britton & Rose.ARGENTINA.ProvMendoza: Dept. Las Heras,04.I.2002,Leuenberger or4793 (SI), PALCTES 7052.Echinopsis ancistrophora Speg.ARGENTINA.ProvSkip: Dept. Church02.II.1995,Kiesling8742 (SI), PAL-CTES 7054.Echinopsis aureaBritton & Rose.ARGENTINA.ProvLa Rioja: Dept. General Ocampo,11.XI.1999,Biurrun et al.4932 (SI), PAL-CTES 7055;Dept.RV Penaloza; 11.XI.1999,Biurrun et al.4933 (SI)-PAL CTES 7364.Echinopsis leucantha(Gillies ex Salm-Dyck) Walp.ARGENTINA.ProvBuenos Aires:LocCardinal Lagliero undated. Kiesling et al1032 (SI) PAL-CTES 7056;ProvCatamarca: Dept. Chamical,undatedKiesling190 (SI), PAL-CTES 7402.Echinopsis mamillosaGurke.ARGENTINA.ProvSalta,undatedCrdenas5022 (LIL), PALCTES 7446.BOLIVIA.Dept.Tarija:Prov.Acinetobacter Arce, Becket al.22150 (SI) PAL-CTES 7057.Echinopsis oxygona(Link) Zucc.ARGENTINA.ProvEntre Ros: Dept. Columbus:El Palmar National Park, La Glorieta, undated,Bacigalupo et al.1603 (SI) PAL-CTES 7448.Echinopsis rhodotrichaSchum.ARGENTINA.ProvFormosa, 18.XI.1999,Di Giacomo433 (SELV) CTES PAL-7058,Dept.Gualeguaych, 23.XI.1999,. Bacigalupo et al. 1793 (SI) 705 PAL-CTEStubiflora Echinopsis(Pfeiff.) Zucc.ARGENTINA.ProvTucumn: Dept. Trancas,03.XII.2003,Leuenberger et al.4903 (LIL) PAL-CTES 7447.Monvillea cavendischii(Monv.) Britton & Rose.ARGENTINA.ProvCorrientes: Dept. Capital,24.IX.1972,Schinini5395 (SELV), PAL-CTES 7256.ProvFormosa: Dept. Lahis,10.XII.1999,Di Giacomo440 (SELV), PAL-CTES 7409;Dept.Patino,20.II.2004,Tressens SG6922 (SI) PALCTES 7410.ProvMissions: DeptSan Ignacio., 23.III.1998, Zuloaga, 6603 (SI), PAL-CTES 7255.Monvillea spegazzini(FAC Weber) Britton & Rose.ARGENTINA.ProvFormosa:Dept.Patino,11.XII.1979,Schinini19597 (SI), PAL-CTES 7257.Dept.Laish,18-II-2000,Di Giacomo452 (SELV), PAL-CTES 7411.

RESULTSGeneral DescriptionAccording to the pollen morphology, are studied gender estenopalnicos, the general characteristics of pollen studied are circular to subcircular prolate spheroidal grains, isopolares, radiosimtricos, medium to large size (48 to 87.5 m?) Field.Tricolpate.Colpi of 15-63 m in length?.Sensed Exina 1-3? M thick, 0.5-2 sexine? Nexin 0.5-1 m? M.SEM confirmed that the tectum is perforated with holes surrounded with annular thickenings, alternating with the same spinules, and nanoespinas microespinas observed with higher and lower density (Table 1).Palynological Description of speciesCereus aethiops(Fig.1AF)large grains, prolate P = 70 (75) 80? m, E = 50 (55) 60? m, P / E = 1.36, subcircular area.Tricolpate.Colposcopy lengths of 50-63? M in length, leaving reduced approximately 13.5? M apocolpios.Drilled, in the same positive and negative elements tectum minors 1IM, making measurement difficult observed with MO.Exina 2? M thick.Sexine nexin 1.4 m 0.6 m?.SEM: The observation allows checking that the perforations and thickening the tectum 0.3 and 0.1 m measured respectively and the measured microespinas supratectales 1m length? varying the distance between them of 2-5 m?.Geographical distribution:Buenos Aires, Catamarca, Crdoba, La Pampa, La Rioja, Mendoza, Rio Black, Salta, San Juan, San Luis, Tucumn.

Fig.1.Cereus aethiopsA: SEM,polar overview.B:. equatorial overviewC:. sculpture detailD:. detailed structureE: MO, equatorial overview.F:. polar overviewCereus argentinensisG: SEM, polar overview.H:equatorial overview.I:detail of sculpture.J: MO, equatorial overview.K:. polar overviewCereus forbesiiL: MEB, polar overview.M:detail of sculpture.N: MO,polar overview.O:equatorial overview.The scale is 31.4 m?A, 33.3 m in?B, 2.6 m in?C, 4 m in?D, 32 m in?E-F;? 28 m inGandK, 28 75 m in?H-J, 2 m in?I, 26 m in?L, 1.5 m in?M, 32 m in?NOT.Cereus argentinensis(Fig. 1GK)large grains, spheroidal, prolate-spheroidal, P = 55 (57.5) 60? m, E = 50 (52) 55? m, P / E = 1.10, a circular area.Tricolpate.Long colposcopy, 50-53? M long, reaching almost to both poles apocolpios approximately 10? M.Tectum drilled on it, very small positive elements arranged irregularly, whose shape can not be determined with the observed MO.Exina 3? M thick.Sexine nexin 2 m 1 m?.MEB: This observation has determined that the tectum perforations from 0.17 to 0.3 m in diameter surrounded by a small annular thickening 0,15 to 0,18 m only? and positive elements are supratectales microespinas measuring 1 m in height, varying the distance between the same 4-6 m?.Geographical distribution:Corrientes and Entre Rios.Cereus forbesii(Fig.1LO)large grains, spheroidal, prolate-spheroidal, P = 55 (57.5) 60? m, E = 50 (52) 55? m, P / E = 1.10, a circular area.Tricolpate.Long colposcopy, 45-50? M long, reaching almost to both poles apocolpios 10? M.Perforated tectum and granular.These granules are very small to be measured with the MO.Exina 2? M thick.Sexine and nexin 1IM.MEB: This observation determines that the tectum is perforated with small holes surrounded by an annular thickening, distinguishable only with high magnification (5000 x) both drilling and thickening measured 0.2 m respectively and?? granule?were observed with MO are nanoespinas supratectales measuring 0.4 m in height, varying the distance of 2-6 m between them?.Geographical distribution:Catamarca, Chaco, Crdoba, Formosa, Jujuy, Tucumn, La Rioja, Salta , Santa FeCereus haenkeanus(Fig. 2AE) large grains, spheroidal, subprolatos P = 55 (56) 57? m E = 40 (43) 47? m, P / E = 1.3, a circular area.Tricolpate.Colpi long, of 40? Meters long, reaching almost to both poles, apocolpios 10? M.Tectum perforations alternating with positive elements whose shape can not be determined with MO.Exina 3? M thick.Sexine nexin 2 m?1 m?.MEB: The observation has determined that 0.3 micro-m in diameter surrounded by a small annular thickening of 1.6 m thick, distinguishable only with high magnification (5000x)? and that the positive elements are supratectales microespinas 1 m high, the distance between them of approximately 3-5 m?.Geographical distribution:Catamarca, Jujuy, Salta.

Fig.2.Cereus haenkeanusA: SEM,polar overview.B:overview equatorial two colpi are observed.C:overview equatorial a colpo is observed.D: MO, polar overview.E:equatorial overview.Cereus stenogonusF: SEM, polar overview.G:equatorial overview.H:detail of sculpture and a colpo partially observed.I: MO, equatorial overview.J:polar overview.Cereus uruguayanusK: MEB, polar overview.L:equatorial overview.M:detail of sculpture.N: MO, polar overview.O:equatorial overview.The scale is 21.5 m?AC, 24 m in?D, 28.6 m in?E, 27 m in?F-G;? 6.6 m inH?, 31 m inIJ?; 25 m inKL, 2 m in?M, 25.7 m in?NOT.Cereus stenogonus(Fig. 2FJ)large grains, spheroidal, prolate-spheroidal P = 55 (57.5) 60? m, E = 50 (51.3) 60? m, P / E = 1.07, a circular area .Tricolpate.? Long colposcopy, 45-55 m in length, reaching almost to the two poles; apocolpios between 5 and 10 m?.Perforated tectum, with positive and negative elements smaller than 1 micrometer, whose shape can not be determined with MO.Exina 3? M thick.Sexine nexin 2 m 1 m?.MEB:? observation has determined that 0.3 micro-m in diameter surrounded by a small annular thickening of 0.2 m thick and the positive elements are supratectales microespinas of1? ?. meters high, varying the distance between them 4-6 mGeographical Distribution:Chaco, Corrientes Misiones.Cereus uruguayanus(Fig. 2KO)large grains, spheroidal-spheroidal subprolate, P = 55 (57.5) 60? m, E = 40 (45) 50? m, P / E = 1.27, a circular area.Tricolpate.Colpi long, of 48? Meters long, reaching almost to both poles, apocolpios between 5-10? M.Perforated tectum, under positive and negative elements are observed 1? M whose shape can not be determined with MO.Exina 2? M thick.Nexin sexine 1.5m 0.5m?.MEB: It can be seen surrounded by possessing a small annular thickening microperforations, both of 0.2 m in diameter and thickness respectively and positive elements are microespinas supratectales of ? 0.9 m in height, varying the distance between them of 2-5 m?.Geographical distribution:Buenos Aires, Entre Rios.Cleistocactus baumannii(Fig. 3AE)medium grains, spheroidal, prolate-spheroidal-spheroidal to subprolate, (P) = 40 (42.5) 45? m, E = 30 (35) 40? m, P / E = 1 21, a circular area.Tricolpate.Colpi 20-25? M long apocolpios ranging between 10 and 15? M.Perforated tectum in the same positive and negative elements less than 1 that can not be defined with the observed MO.Exina 1.5? M thick.1IM and nexin sexine of 0.5 m?.MEB:? tectum perforations measured 0.17 m in diameter and are surrounded by annular thickening of 0.22 to 0.28 m and are between 0.81 supratectales microespinas -0.91 m high, varying from 1-4 m distance?.Geographical distribution:Catamarca, Chaco, Crdoba, Corrientes, Entre Ros, Formosa, Jujuy, La Rioja, Salta, Santiago del Estero, Santa Fe, Tucumn .

Fig.3.Cleistocactus baumanniiA: SEM,polar overview.B:. equatorial overviewC:. sculpture detailD: MO, polar overview.E:detail of optical section.Cleistocactus hyalacanthusF: SEM, polar overview .G:equatorial overview.H: MO, polar overview.I:equatorial overview.Cleistocactus samaragdiflorusJ: SEM,polar overview.K:equatorial overview.L:detail of sculpture.M: MO,polar overview .N:equatorial overview.The scale bar represents 35 m?A,BandD,? 5 m inC;? 7.5 m inE;? 23.75 m inFH,? 27 m inI;? 16.25 m inJ; 21 8 m in?K, 3 m in?L, 28.3 m in?MN.Cleistocactus hyalacanthus(Fig.3FI)medium grains, spheroidal, prolate-spheroidal-spheroidal to subprolate, P = 45 (47.5) 50? m, E = 38 (41.5) 45? m P / E = 1 circular field 11.Tricolpate.Colposcopy from 25 to 27.5 ? M length apocolpios 10 to 15? M.Perforated tectum in the same positive and negative elements 1IM minors who fail to distinguish the observed MO.Exina 1.5? M thick.Nexin sexine of 1IM and 0.5 m?.SEM: with perforations of 0.17 m is observed that the tectum is perforated and surrounded by annular thickenings 0.15-0.20 m supratectales nanoespinas of alternating? ? 0.43 to 0.45 m and 0.52 m supratectales microespinas height, varying the distance between them of 2-5 m?.Geographical distribution:Jujuy, Salta.Cleistocactus smaragdiflorus(Fig.3JN)medium grains, spheroidal, prolate-spheroidal-spheroidal to subprolate, P = 40 (42.5) 45? m (E) = 25 (32.5) 40? m, P / E = Circular 1.30 level.Tricolpate.Long colpi 25-30? M in length, 12 to 15 apocolpios? M.Perforated tectum on the same minor positive and negative elements are observed 1? M uniformly distributed.Exina 1.5? M thick.Sexine nexin 1 m 0.5 m?.MEB:? tectum is appreciated with the drilling of 0.14-0.15 m, surrounded by annular thickening of 0.15-0.18 m nanoespinas supratectales 0 ?, 49 m, supratectales microespinas of 0.74-0.80 m in height and the distance between them varies 2-4 m?.Geographical distributionCatamarca.Denmoza rhodacantha(Fig. 4AE)medium grains, spheroidal, prolate-spheroidal-spheroidal to subprolate, P = 38 (44) 50? m, E = 30 (37.5) 45? m, P / E = 1.17 , a circular area.Tricolpate.Colpi 30 to 35? M long, 12 to 17 apocolpios? M.Operculum residues are observed.Perforated tectum and granular.Exina 2? M thick level where Ecuador is increasing gradually to reach in the pole 4? M.Sexine and nexin the same thickness 1 m level Ecuador and 2 ma level poles?.MEB: At high magnification (5000x) can be seen that the tectum is perforated in both the micro and thickening the surrounding measured 1 each 1? m.The granulations are microespinas supratectales 0.8 m tall, irregularly arranged, varying the distance between them of 2-5 m?.Geographical distribution:Catamarca, La Rioja, Mendoza, Salta, San Juan, Tucumn.

Figure 4.Denmoza rhodocanthaA: SEM, polar overview.B:overview equatorial,C:detail of sculpture.D: MO, equatorial overview.E:. polar overviewEchinopsis ancistrophoraF: SEM, polar overview.G:detail of sculpture.H: MO, polar overview.I:equatorial overview.Echinopsis aureaJ: SEM, polar overview.K:detail of sculpture.L: MO, polar overview.M: equatorial overview.Echinopsis leucanthaN: MEB, polar overview.O:detail of sculpture.The scale is 18.75 m in?A-B, 5.3 m in?C, 21.5 m in?DE; 18.2 m 5 m in G, 22.05 m in??H, 25, 6I;? 20.7 m inJ, 4 m in?K, 24.2 m in?L, 29.5 m in?M, 28 m in?NOT.Echinopsis ancistrophora(Fig.4FI)medium grains, spheroidal-spheroidal subprolate, (P) = 45 (47.5) 50? m, (E) = 35 (37.5) 40? m, P / E = 1 26, a circular area.Tricolpate.Long colposcopy of 37.5 to 50? M long, 15 to 25 apocolpios? M.Tectum with small perforations on the same positive and negative elements less than 1? M are observed.Exina 2? M thick.Sexine and nexin 1 m?.MEB:? 0.25 The measured micro-m in diameter, surrounded by annular thickening of from 0.34 to 0.36 m and 1.23 m supratectales spinules high, varying irregularly arranged distance from 1-10 m?.Geographical distribution:Jujuy, Salta, Tucumn.Echinopsis aurea(Fig.4JM)large grains, spheroidal-spheroidal subprolate, (P) = 54 (62) 70? m, E = 40 (42.5) 45? m, P / E = 1.45, field Circular.Tricolpate.Long colposcopy of 32.5 to 50? M length apocolpios 15 to 20? M.Perforated tectum, positive and negative elements 1IM minors are observed.Exina 2? M thick.?. Sexine and nexin 1 mMEB: Distinguished the tectum has microapertures 0.22 to 0.25 m in diameter surrounded annular thickenings measuring from 0.34 to 0.36 m?.? Presence of supratectales microespinas 0.53 to 0.83 m in height, the distance between them is 3-6 m?.Geographical distributionCrdoba.Echinopsis leucantha(Fig.4NO) and (Fig. 5AC)large grains, prolate, (P) = 65 (72.5) 80? m, (E) = 38 (44) 50? m, P / E = Circular 1.64 level.Tricolpate.Colpi 15-25? M in length, 12 to 15 apocolpios? M.Perforated tectum on the same minor positive and negative elements are observed 1? M.Exina 2? M thick.Nexin sexine and 1 m?.SEM: tectum is observed that is perforated with holes of 0.21 to 0.37 m surrounded by annular thickenings measuring 0.16-0.24 m?.? Supratectales microespinas presence of 0.70 to 0.87 m, arranged irregularly varying the distance of 1-7 m?.Geographical Location:Buenos Aires, Catamarca, La Rioja, Mendoza, Rio Black, Salta, Santiago del Estero, San Juan, San Luis, Tucumn.

Figure 5.Echinopsis leucanthaA: MO, polar overview.B:. equatorial overviewC:. cutting opticalEchinopsis mamillosa.D: SEM, polar overview.E:. equatorial overviewF:. sculpture detail ofG: MO, polar overview.H:equatorial overview.Echinopsis oxigona.I: MEB, polar overview.J:detail of sculpture.K: MO, polar view.L:equatorial view.M:optical section.The scale represents 5.8 m in?A, 22.5 m in?B, 42.6 m in?C, 26.6 m in?D, 3.3 m in?E;? 31.5 m inF, 28 m in?G, 9.9 m?H, 22.8 m?I, 3.2 m?J,30 m?K-L, 11.5 m?M.Echinopsis mamillosa(Fig.5DH)large grains, prolate, (P) = 70 (80) 90? m, (E) = 50 (60) 70? m, P / E = 1.33, a circular area.Tricolpate.Colpi 62.5? M in length and not become attached to the poles, apocolpios 15 to 30? M.Tectum granulated on the same minor positive and negative elements are observed 1IM, Exina 1.5? M thick.Sexine nexin 1 m and 0.5 m?.SEM: tectum is observed that is perforated with holes of from 0.21 to 0.59 m surrounded by annular thickenings measuring 0.25-0.45 m?.? Presence of supratectales microespinas measuring 0.99 m in height, arranged irregularly varying the distance 3-9 m?.Geographical distributionSalta.Echinopsis oxygona(Fig.5IM)large grains, spheroidal, prolate-spheroidal to prolate, (P) = 55 (60) 65? m, E = 45 (51.5) 58? m, P / E = 1.16 , a circular area.Tricolpate.Colposcopy lengths of 27.5 to 37.5? M length apocolpios 15? M.Perforated tectum, under positive and negative elements are observed 1? M not differ in MO.Exina 2? M thick.? Sexine and nexin 1 m.MEB: It is confirmed that the tectum is perforated with holes measuring from 0.29 to 0.38 m in diameter, surrounded by an annular thickening of 0.27 m supratectales spinules of 1? ? 15 m and the distance between them is 3-8 m?.Geographical distribution:Buenos Aires, Entre Rios and Santa FeEchinopsis rhodotricha(Fig.6AE)large grains, prolate, (P) = 72 (78.5) 85? m, (E) = 35 (42.5) 50? m, P / E = 1.84, field Circular.Tricolpate.Colpi 25-30? M, apocolpios 15 to 20? M.Granular tectum on the same minor positive and negative elements are observed 1? M.Exina 2? M thick.Nexin sexine and 1 m?.SEM: tectum is observed that is perforated with perforations of between 0.25-0.27 m surrounded by annular thickenings of 0.21 to 0.22 m?.Presence of supratectales microespinas 0.86 m of 1.11 to 1.31 m supratectales spinules, arranged irregularly, varying the distance between 3-7 m??.Geographical distribution:Chaco, Corrientes, Entre Ros, Formosa, Santiago del Estero, Santa Fe

Fig.6.Echinopsis rhodotrichaA: SEM,polar overview.B:. sculpture detail ofC:. detailed structureD: MO, polar overview.E:. equatorial overviewEchinopsis tubifloraF: SEM, equatorial overview.G:detail of sculpture.H: MO, polar overview.I:equatorial overview.The scale is 20.2 m?A;? 4.3 m inB;? 5.2 m inC, 25 m in?D;? 34.1 m inE, 28.3 m in?F; ? 4 m inG, 30 m in?HandI.Echinopsis tubiflora(Fig.6FI)large grains, prolate, (P) = 70 (77.5) 85? m, (E) = 45 (50) 55? m, P / E = 1.12, a circular area.Tricolpate.Colpi 50-60? M in length, 20 to 25 apocolpios? M.Granular tectum on the same minor positive and negative elements are observed.Exina 2? M thick.Sexine and 1IM nexin.SEM: tectum is observed that is perforated with holes measuring between 0.35-0.47 m surrounded by annular thickening ranging between 0.22 to 0.27 m?.? Presence of spinules measuring 1.08 to 1.41 m, the distance between them is 2-10 m?Geographical Distribution:Salta, Tucumn.Monvillea cavendischii(Fig.7AE)large grains, spheroidal, prolate-spheroidal-spheroidal to subprolate, (P) = 55 (57.5) 60? m, (E) = 35 (45) 55? m, P / E = 1.27, a circular area.Tricolpate.Long colpi 26-37? M in length, 15 to 17 apocolpios? M.Granular tectum in the same minor positive and negative elements are observed 1? M, arranged regularly.Exina 1? M thick.Nexin sexine and 0.5 m?.SEM: tectum is observed that is perforated with perforations surrounded 0.25-0.30 m annular thickenings measuring 0.28m?.? Presence of spinules supratectales 1.15 m tall, regularly arranged and the distance between them is 1-5 m?.Geographical distribution:Chaco, Corrientes, Entre Ros.Formosa, Jujuy, Salta.

Fig.7.Monvillea cavendischiiA: SEM,polar overview.B:. equatorial overviewC:. sculpture detailD: MO, polar overview.E:. equatorial overviewMonvillea spegazziniiF: SEM, polar overview.G:detail of sculpture.H: MO, polar overview.I:equatorial overview.The scale represents 27.5 m?A,B,DandE, 5 m in?C, 20 m?F;? 2 m inG,? 24.8 m inHandI. Monvillea spegazzinii(Fig.7FI)large grains, prolate, P = 65 (62.5) 75? m, E = 35 (37.5) 40? m, P / E = 1.6, a circular area.Tricolpate.Long colpi 35-45? M long, 20 apocolpios? M.Drilled in the same tectum under positive and negative elements are observed 1? M regularly arranged.Exina 1? M thick.Nexin sexine and 0.5 m?.SEM: tectum is observed that is perforated with holes of 0.24 to 0.30 m surrounded by annular thickenings measuring from 0.25 to 0.28 m?.? Presence microespinas measuring 0.90 m high and 1.20 m spinules high, regularly arranged and the distance between them is 1-6 m?.Geographical distribution:Chaco, Formosa.DISCUSSION AND CONCLUSIONSThe palynological analysis of the genera studied, not only provides data on each of the Argentine species, but extends previous studies by other authors for Cactaceae pollen.According Nowicke work (1975), who studied 12 species of belonging to the Cactaceae subfamily Opuntoideae there characters that are common to most members of the family. spinulose tectum and finding tricolpate drilled beads, and pantoporados pantocolpadosMeanwhile, Leuenberger (1976) studied the pollen of 600 species of Cactaceae from different parts of America, of which six of them:Cereus forbesii,Cleistocactus baumannii,C.smaragdiflorus,Denmoza rhodocantha, Echinopsis rhodotricha, Monvillea cavendischiiandM.spegazziniiare studied here.Most of the characters provided by the author match what I studied in this paper, except the number of openings of some species.The same author when studyingCleistocactus baumannii,C.smaragdiflorus,Denmoza rhodocanthaandrhodotricha Echinopsisfound that these species had tricolpate most grains and a smaller percentage (1-2%) pantocolpados grains.Pollen grains of the same species studied here are all tricolpate.Likewise, the author analyze the pollenCereus forbesii,Monvillea cavendischiiandM.spegazziniitricolpate are observed, which is consistent with the analysis in these species.This difference suggests the possibility of pollen used hybrids, having worked in some cases grown material.Garrallaet al.(2008) analyzed pollen Argentine speciesRebutiaS.str.and recognized two groups, based on the type of opening: tricolpate and pantocolpados.Also another character that was taken into account the shape of the pollen grains: suboblatos a prolate spheroidal and subprolatos or.pollen study of Argentine species ofRhipsalisGaerth.conducted by Salgado & Garralla (2008), allowed us to observe that the main differences between taxa studied refer to the number of openings and the arrangement of the same on the surface of the pollen grains.Established 2 groups. 3-colpados and pantocolpados latter with 6 and 8 respectively colpiLater, Square & Garralla (2009) studied the pollen grains of the species belonging to the Argentine and Pereskoideae Maihuenioideae subfamilies.The authors found no differences in the number and arrangement of openings establishing two groups: tricolpate pantocolpados grains and beans.Among tricolpate are species of the genusMaihuenia(FAC Weber) K. Schum.(M. patagonicaandM. poepigii) who in turn can be separated by the shape and size of grains and length of the colpi.Instead between pantocolpados are speciesPereskiaMill, where colposcopy were found in number of 6-14.Comparing pollenCereusMill,CleistocactusLem.,DenmozaBritton & Rose,EchinopsisZucc.andMonvilleaRose Britton, have close similarity with these speciesMaihueniain the number of openings in the form and sculpture of the grains.The genera referred to above present tricolpate grains prolate spheroidal or sensed and exine perforations surrounded with annular thickenings, with spinules, or nanoespinas microespinas, these data are consistent with other, already published (Kurtz, 1963; Leuenberger, 1976.1997; & Garralla Square, ., 2009)Finally, the palynological of genres Cactaceae-Cactoideae studied, characters reveal that there is much affinity Pereskioideae and Maihuenoideae and not with Opuntioideae as they had mentioned (Leuenbenger, 1976, 1997; Garralla & Square, 2009) .According to palynological analysis in this work we can conclude the following:* Due to the homogeneity of the species studied, it was only possible to separate it into two types: TypeCereus argentinensisand TypeEchinopsis mamillosa, given shape and size of the . pollen grains* next key palynofloral according to the characteristics observed in both types is provided:Key to the pollen types1.40-65 spheroidal? M diameter..............TypeCereus argentinensis1.?Prolate 70 to 90? M diameter..............TypeEchinopsis mamillosaThe typeCereus argentinensispresented inCereus forbesii, C. haenkeanus, C. stenogonus, Cleistocactus hyalacanthus, C. smaragdiflorus,Denmoza rhodocantha,Echinopsis ancistrophora, E. aurea, E. oxygonaandMonvillea cavendischii.TypeEchinopsis mamillosapresented inCereus aethiops, Echinopsis leucantha, E. rhodotricha, E. tubifloraandMonvillea spegazzinii.AcknowledgementsWe are especially grateful to the Missouri Botanical Garden Institute (SI), Northeast Institute of Botany (SELV), Instituto Miguel Lillo (LIL) for the material given.Dr. Beat Leuenberger for the material provided.Dr. Roberto Kiesling for critically reading the manuscript and St. Eva Acevedo (Laboratory Technician CONICET) for the preparation of the pollen material.Serving Electron Microscopy (SEM), the Ministry of Science and Technology, National University of the Northeast.REFERENCES1.SQUARE, GA & SS GARRALLA.2009.Palynology of the generaMaihuenia(Maihuenioideae) andPereskia(Pereskioideae), Cactaceae, of Argentina.Bonplandia18 (1) :5-12.[Links]Two.Erdtman, G. 1966.Pollen morphology and plant taxonomy.Angiosperms.New York: Hafner Publ.Co. [Links]Three.GARRALLA, SS & GA SQUARE.2007.Pollen morphology of the Cactaceae of Argentina (Opuntoideae)AustrocylindropuntiaBackeb,.MaihueniopsisSpeg,.OpuntiaMillandTephrocactusLem.Palaeobotany and Palynology Review of146: 1-17.[Links]April.GARRALLA, SS, SQUARE NB MURUAGA & GA.2008.Pollen morphology of Argentine speciesRebutiaS.str.(Cactaceae, Cactoideae).Darwiniana46 (2): 270-278.[Links]May.KIESLING, R. 2005.Cactaceae.In NM Bacigalupo (Director), Illustrated Flora of Entre Rios (Argentina) IV: 258-265.[Links]6.KIESLING, R., L., LARROCA, J. FANDEZ, D. FANDEZ, D. & S. Metzing ALBESIANO (Wigginsia).2008.Cactaceae.In FO Zuolaga, O. Morrone & MJ Belgrano (eds.), Catalogue of the Vascular Plants of the Southern Cone.Monogr.Syst.Bot.Gard.107: 1711-1830[Links]

7.KURTZ, EB1963.Pollen morphology of the Cactaceae,Grana Palynol.IV (3) :367-372.[Links]8.LEUENBERGER, BE 1976.Die Morphologie der Cactaceae pollen Dissertationes Botanicae, 1-321.[Links]9.LEUENBERGER, BE 1997.Mahuenia, monograph of Patagonian genus of Cactaceae.Bot.Jarb.Syst.119:1-92.[Links]10.Nilsson, S. & J. PRAGLOWSKI.1992.Erdtman?Handbook of Palynology, 2 da.Ed Munksgaard, Copenhagen.[Links]11.NOWICKE, JK 1975.Pollen morphology in the order Centrospermae.Grana Palynol.15: 51-77.[Links]12.PUNT, W., PP HOEN, S. & A. LE THOMAS BLACKMORE.2007.Glossary of pollen spore terminology.Review of Paleobotany and Palynology143: 1-81.[Links]13.SALGADO, CR & SS GARRALLA.2008.Xixo Meeting of Scientific and Technical Communications and Outreach Meeting.Pollen Argentine species of the genusRhipsalisGaertn.(Cactaceae-Cactoideae).FCA-UNNE.[Links]