Bulletin of the Botanical Society of Argentina

On-line version ISSN 1851-2372

BullSocArgent.Bot.Crdobav.45n.12Jan./June.2010

Paleobotany And PALYNOLOGY

Palynological studies of Argentinean species of the

genera Cereus , Cleistocactus , Denmoza , Echinopsis andMonv
illea (Cactaceae, Cactoideae)

Elsa LATTAR 1 , 2 & Graciela Square 1 , 3

1

Chair of Palynology, Faculty of Natural Sciences and Surveying, Universidad Nacional del
Nordeste, Corrientes, Argentina. E-mail: elsalattar@yahoo.com.ar ;elsilattar@gmail.com
2
Northeast Institute of Botany, IBONE
3
Center for Applied Coastal Ecology, CECOAL

Abstract : The pollen morphology of 19 species belonging to five genera

(studied Cereus , Cleistocactus , Denmoza , Echinopsisand Monvillea ) of Cactoideae
(Cactaceae) in Argentina subfamily were described using light microscopy and scanning
electron microscopy: Cereus aethiops Haw ., C. argentinensis Britton &
Rose, C. forbesii CFFrst Otto ex., C. haenkeanus FAC Weber ex K.
Schum., C. stenogonus K. Schum., C. uruguayanus R. 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. aurea Britton & Rose, E. leucantha (Gillies ex Salm-Dyck)
Walp., E. mamillosa Gurke, E.oxygona (Lynk)
Zucc., E. rhodotricha K. Schum., E. tubiflora (Pfeiff. & Zucc.) Monvillea cavendischii (Monv.)
Britton & Rose and M.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 determined Cereus argentinensis and
Pollen Exchange mamillosa 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
generates Cereus , Cleistocactus , Denmoza , Echinopsis andMonvillea (Cactaceae,
Cactoideae) . Pollen grains morphology of nineteen species generates Belonging to five
( Cereus ,Cleistocactus , Denmoza , Echinopsis and Monvillea ) of the subfamily Cactoideae
(Cactaceae) in Argentina, are Described using optical and scanning electron
microscopy: Cereus aethiops Haw. C. argentinensis Britton & Rose, C. forbesii CFFrst Otto
ex., C.haenkeanus FAC Weber ex K.
Schum., C. stenogonus K. Schum., C. uruguayanus R. 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. aurea Britton & Rose, E. leucantha (Gillies ex Salm-Dyck)
Walp., E. mamillosaGurke, E. oxygona (Lynk)
Zucc., E. rhodotricha K. Schum., E. tubiflora (Pfeiff. & Zucc.) and Monvillea
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 type Cereus
argentinensis and pollen type mamillosa Echinopsis .A key to identify identity and Distinguish
the pollen types was given.
Key words: Pollen; Morphology; Cactaceae; Taxonomy.

INTRODUCTION
Cactaceae 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 Kiesling et 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 , Echinopsis and Monvillea .
publications referred pollen Cacti are scarce in Argentina. Internationally can
cite the work of Kurtz (1963), who analyzed the pollen of some species
of Cleistocactus , Denmoza and Echinopsis, 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
generaAustrocylindropuntia Backeb, Maihueniopsis .
Speg, Opuntia Mill and Tephrocactus Lem. (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. Garralla et al . (2008) analyzed the pollen morphology of the
species Rebutia K. Schum., Mainly finding differences in the type of
openings. Then Garralla & Salgado (2008) studied pollen
species Rhipsalis Gaertn. 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
published Maihueniodeae and Pereskioideae of 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 METHODS
The 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. For Cleistocactus yalacanthus only 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 Punt et al. (2007), for the name of the spines according to their
different heights.
Material examined
Cereus aethiops Haw. ARGENTINA. Prov Skip: Dept. General
Cafayate, 04.I.1972, Krapovickas et al. 20621 (SELV), PAL-CTES
7019; Prov Corrientes: Dept. Capital , 18.XII.2003, Meza Torres 163 (SELV),
PAL-CTES 7403. Cereus argentinensis Britton &
Rose. ARGENTINA. Prov Corrientes: Dept. Lavalle, 23.XII.1979,Schinini et
al. 19064 (SI), PAL-CTES 7020. Cereus forbesii Frst CF Otto
ex. ARGENTINA. Prov Formosa: Dept Laish, 10.XII.1999, Di Giacomo 438
(SELV), PAL-CTES 7407. Prov Salta, Dept.
Rivadavia,03.III.1984, Schinini, 19583 (SELV), PAL-CTES 7406. Cereus
haenkeanus FAC Weber ex Schum.ARGENTINA. Prov Jujuy. Dept. San
Pedro , San Pedro to Santa Clara, 11.XI.1995, Kiesling et al. 8719 (SELV),

PAL-CTES 7413. Cereus

stenogonus K. Schum. ARGENTINA. Prov Corrientes : Dept . Capital,25.X.20
03, Meza Torres . 161 (SELV), PAL-CTES 6808 Cereus
uruguayanus R. Kiesling. ARGENTINA.Prov Corrientes: Dept.
Capital, 12.XII.2003, Meza Torres 164 (SELV), PAL-CTES
7305. Cleistocactus baumannii (Lem.) Lem. ARGENTINA. Prov Entre Ros:
Dept. La Paz, Step Yunke, 03.XI.1998, . Bacigalupo et al 1666 (SI), PALCTES 7022; Prov Formosa: Dept. Laish, 18.XI.1999, Di Giacomo 432
(SELV), PAL-CTES 7408. BOLIVIA. Dept. Tarija , 20.XII.1994, Zuloaga et
al . 1365 (SI), PAL-CTES 7024. Cleistocactus hyalacanthus (K. Schum.)
Gosselin. ARGENTINA, Prov Jujuy . Schwerdtfeger 17649 (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.Prov Catamarca: Dept.
Capital, undated Ulibarri 895 (SI), PALCTES 7025. Denmoza
rhodacantha (Salm-Dyck) Britton & Rose. ARGENTINA. Prov Mendoza: Dept.
Las Heras, 04.I.2002, Leuenberger or 4793 (SI), PALCTES 7052. Echinopsis
ancistrophora Speg. ARGENTINA. Prov Skip: Dept.
Church 02.II.1995, Kiesling8742 (SI), PAL-CTES 7054. Echinopsis
aurea Britton & Rose. ARGENTINA. Prov La 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: Loc Cardinal Lagliero undated . Kiesling et al 1032 (SI) PAL-CTES
7056; Prov Catamarca: Dept. Chamical, undated Kiesling 190 (SI), PAL-CTES
7402. Echinopsis
mamillosa Gurke. ARGENTINA.Prov Salta, undated Crdenas 5022 (LIL),
PALCTES 7446. BOLIVIA. Dept. Tarija: Prov . Acinetobacter Arce, Beck et
al. 22150 (SI) PAL-CTES 7057. Echinopsis oxygona (Link)
Zucc. ARGENTINA. Prov Entre Ros: Dept. Columbus: El Palmar National
Park, La Glorieta, undated, Bacigalupo et al. 1603 (SI) PAL-CTES
7448. Echinopsis rhodotricha Schum. ARGENTINA. Prov Formosa ,
18.XI.1999, Di Giacomo 433 (SELV) CTES PAL-7058, Dept. Gualeguaych,
23.XI.1999, . Bacigalupo et al . 1793 (SI) 705 PAL-CTES tubiflora
Echinopsis (Pfeiff.) Zucc. ARGENTINA. Prov Tucumn: Dept.
Trancas, 03. XII.2003, Leuenberger et al. 4903 (LIL) PAL-CTES
7447. Monvillea cavendischii (Monv.) Britton &
Rose. ARGENTINA. Prov Corrientes: Dept. Capital, 24.IX.1972, Schinini 5395
(SELV), PAL-CTES 7256. Prov Formosa: Dept. Lahis, 10.XII.1999, Di
Giacomo 440 (SELV), PAL-CTES 7409; Dept. Patino, 20.II.2004, Tressens
SG 6922 (SI) PALCTES 7410.Prov Missions : Dept San Ignacio. ,
23. III. 1998, Zuloaga, 6603 (SI), PAL-CTES 7255. Monvillea spegazzini(FAC
Weber) Britton &
Rose. ARGENTINA. Prov Formosa : Dept. Patino, 11.XII.1979, Schinini 1959
7 (SI), PAL-CTES 7257. Dept. Laish, 18-II-2000, Di Giacomo 452 (SELV),
PAL-CTES 7411.

RESULTS
General Description
According 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 species
Cereus aethiops ( Fig.1 AF )
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 aethiops A: SEM, polar overview. B: . equatorial overview C: . sculpture

detail D: . detailed structure E: MO , equatorial overview. F: . polar overview Cereus
argentinensis G: SEM , polar overview. H: equatorial overview. I: detail of sculpture. J: MO ,
equatorial overview. K: . polar overview Cereus forbesii L: 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 in G and K , 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. 1 GK )

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 . 1 LO )
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 Fe
Cereus haenkeanus ( Fig. 2 AE ) 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 haenkeanus A: 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 stenogonus F: SEM , polar overview. G: equatorial overview. H:detail of
sculpture and a colpo partially observed. I: MO , equatorial overview. J: polar
overview. Cereus uruguayanus K: 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 in H ?, 31 m in IJ ?; 25 m in KL , 2 m in? M ,
25.7 m in? NOT .

Cereus stenogonus ( Fig. 2 FJ )

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 m Geographical Distribution: Chaco, Corrientes Misiones.
Cereus uruguayanus ( Fig. 2 KO )
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. 3 AE )
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 baumannii A: SEM, polar overview. B: . equatorial overview C: .

sculpture detail D: MO , polar overview. E:detail of optical section. Cleistocactus
hyalacanthus F: SEM , polar overview . G: equatorial overview. H: MO , polar
overview. I:equatorial overview. Cleistocactus samaragdiflorus J: SEM, polar
overview. K: equatorial overview. L: detail of sculpture. M: MO,polar overview . N: equatorial
overview. The scale bar represents 35 m? A , B and D ,? 5 m in C ;? 7.5 m in E ;? 23.75 m
in FH ,? 27 m in I ;? 16.25 m in J ; 21 8 m in? K , 3 m in? L , 28.3 m in? MN .

Cleistocactus hyalacanthus ( Fig.3 FI )

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.3 JN )
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 distribution Catamarca.
Denmoza rhodacantha ( Fig. 4 AE )
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 rhodocantha A: SEM , polar overview. B: overview equatorial, C: detail

of sculpture. D: MO , equatorial overview. E: . polar overview Echinopsis ancistrophora F:
SEM , polar overview. G: detail of sculpture. H: MO , polar overview. I:equatorial
overview. Echinopsis aurea J: SEM , polar overview. K: detail of sculpture. L: MO , polar
overview. M: equatorial overview.Echinopsis leucantha N: 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, 6 I ;? 20.7 m in J , 4 m in? K , 24.2 m in? L , 29.5 m in? M , 28 m
in? NOT .

Echinopsis ancistrophora ( Fig.4 FI )

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.4 JM )
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 m
MEB: 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 distribution Crdoba.
Echinopsis leucantha ( Fig.4 NO ) and ( Fig. 5 AC )
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 leucantha A: MO , polar overview. B: . equatorial overview C: . cutting

optical Echinopsis mamillosa. D: SEM, polar overview. E: . equatorial overview F: . sculpture
detail of G: 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 in F , 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.5 DH )

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 distribution Salta.
Echinopsis oxygona ( Fig.5 IM )
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 Fe
Echinopsis rhodotricha ( Fig.6 AE )
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 0.250.27 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 rhodotricha A: SEM, polar overview. B: . sculpture detail of C: . detailed

structure D: MO , polar overview. E: . equatorial overview Echinopsis tubiflora F: SEM ,
equatorial overview. G: detail of sculpture. H: MO , polar overview. I: equatorial overview. The
scale is 20.2 m? A ;? 4.3 m in B ;? 5.2 m in C , 25 m in? D ;? 34.1 m in E , 28.3 m in? F ; ? 4
m in G , 30 m in? Hand I .

Echinopsis tubiflora ( Fig.6 FI )

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.7 AE )
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 0.250.30 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 cavendischii A: SEM, polar overview. B: . equatorial overview C: . sculpture

detail D: MO , polar overview. E: . equatorial overview Monvillea spegazzinii F: SEM , polar
overview. G: detail of sculpture. H: MO , polar overview. I: equatorial overview. The scale
represents 27.5 m? A , B , D and E , 5 m in? C , 20 m? F ;? 2 m in G ,? 24.8 m in H and I .

Monvillea spegazzinii ( Fig.7 FI )

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 CONCLUSIONS
The 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 pantocolpados
Meanwhile, 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
cavendischii and M. spegazzinii are 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 studying Cleistocactus
baumannii , C. smaragdiflorus , Denmoza rhodocantha and rhodotricha
Echinopsis found 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 pollen Cereus
forbesii , Monvillea cavendischii and M. spegazzinii tricolpate 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.
Garralla et al. (2008) analyzed pollen Argentine species Rebutia S. 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 of Rhipsalis Gaerth. 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 colpi
Later, 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 genus Maihuenia (FAC Weber) K.
Schum. ( M. patagonica and M. poepigii ) who in turn can be separated by the
shape and size of grains and length of the colpi. Instead between
pantocolpados are speciesPereskia Mill, where colposcopy were found in
number of 6-14. Comparing
pollen Cereus Mill, CleistocactusLem., Denmoza Britton &
Rose, Echinopsis Zucc. and Monvillea Rose Britton, have close similarity with

these species Maihuenia in 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 argentinensis and Type Echinopsis
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 types
1. 40-65 spheroidal? M diameter
. . . . . . . . . . . . . . Type Cereus argentinensis
1.? Prolate 70 to 90? M diameter
. . . . . . . . . . . . . . Type Echinopsis mamillosa
The type Cereus argentinensis presented in Cereus forbesii, C. haenkeanus,
C. stenogonus, Cleistocactus hyalacanthus, C. smaragdiflorus , Denmoza
rhodocantha , Echinopsis ancistrophora, E. aurea, E. oxygonaand Monvillea
cavendischii.
Type Echinopsis mamillosa presented in Cereus aethiops, Echinopsis
leucantha, E. rhodotricha, E. tubifloraand Monvillea spegazzinii.
Acknowledgements
We 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.
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