Pollen and SPoreS morPhology from the holocene

of the Iberá wetlandS In northeaStern argentIna

morfología del Polen y eSPoraS del holoceno de loS eSteroS del

Iberá en el noreSte de argentIna

1

2

Lionel Fernandez Pacella * & Mercedes Di Pasquo

Summary

Background and aims: Taxonomic works of the modern pollen grains from the

vegetation of the Iberá Wetlands from northeastern Argentina, and other areas of

the Corrientes province, have mostly been carried out in last decade. However, very

few of these taxonomic works include illustration of palynomorphs. The objective

of this contribution is to provide the ﬁrst morphological records of palynomorphs of

angiosperms, ferns, lycophytes and bryophytes from sediments from the Holocene

of the Iberá Wetlands.

M&M: Core samples obtained by mean of a Livingston-type sampler from six lakes in

their central and deepest parts on the western margin of the Iberá Wetlands were

analyzed.

1

. Centro de Ecología Aplicada del

Litoral (CECOAL-CONICET-UNNE),

Dpto. Biología-FaCENA-UNNE, Ruta

5

2

, Km 2,5, Corrientes, Argentina.

. Laboratorio de Palinoestratigrafía

y Paleobotánica, CICyTTP-CONICET,

Dr. Materi y España s.n., Diamante,

Entre Ríos, Argentina. medipa@

cicyttp.org.ar

Results: Fifty-ﬁve types of palynomorphs are described and illustrated: 46 pollen

types correspond to 27 families of angiosperms and nine trilete spore-types to ferns,

lycophytes and bryophytes. Information to diﬀerentiate local (species that form part of

the natural vegetation of the Iberá Wetlands) and extra-local taxa (those that do not

belong to the Iberá), that achieved to depocenters mostly by means of wind currents

coming from farther regional vegetation, was included.

Conclusions: Pollen grains and spores identiﬁcations up to species level enhances

paleoenvironmental reconstructions based on more accurate ecologic information

and geographical distribution. This work broadens the knowledge of the palynological

ﬂora of northeastern Argentina and it will contribute to diﬀerentiate the local vegetation

from the extra-local in future paleocological and paleoenvironmental interpretations.

*

lionelpacella@yahoo.com.ar

Citar este artículo

FERNANDEZ PACELLA, L. & M. DI

PASQUO. 2022. Pollen and spores

morphology from the Holocene of

the Iberá Wetlands in northeastern

Argentina. Bol. Soc. Argent. Bot. 57:

727-750.

DOI: https://doi.

org/10.31055/1851.2372.v57.

n4.35662

KeywordS

Corrientes Province, hydrographic system Iberá, holocene, palynomoph.

reSumen

Introducción y objetivos: Los trabajos taxonómicos de polen actual de la vegetación

de los esteros del Iberá en el noreste de Argentina y otras áreas de la provincia

de Corrientes se han llevaron a cabo mayoritariamente en la última década. Sin

embargo, muy pocos de estos trabajos incluyen la ilustración de los palinomorfos.

El objetivo de esta contribución es proporcionar el primer registro morfológico de

palinomorfos de angiospermas, helechos, licoﬁtas y brioﬁtas de sedimentos del

Holoceno de los Esteros del Iberá.

M&M: Se analizaron muestras de núcleos obtenidos por medio de un muestreador

tipo Livingston de seis lagos en sus partes central y más profunda en el margen

occidental de los Esteros del Iberá.

Resultados: Se describen e ilustran 55 tipos de palinomorfos: 46 tipos de polen

corresponden a 27 familias de angiospermas y nueve tipos de esporas triletas

a helechos, licoﬁtas y brioﬁtas. Se incluye información para diferenciar taxones

locales (que forman parte de la vegetación natural de los Esteros del Iberá) y extra-

locales (aquellos que no pertenecen al Iberá) que lograron llegar a los depocentros

provenientes de vegetación regional más lejana.

Conclusiones: La identiﬁcación de granos de polen y esporas hasta el nivel de especie

mejora las reconstrucciones paleoambientales basadas sobre información ecológica

y distribución geográﬁca más precisas. Este trabajo amplía el conocimiento de la ﬂora

palinológica del noreste de Argentina y contribuirá a diferenciar la vegetación local de

la extra-local en futuras interpretaciones paleoecológicas y paleoambientales.

Recibido: 20 Nov 2021

Aceptado: 19 Ago 2022

Publicado impreso: 30 Dic 2022

Editor: Gonzalo Márquez

PalabraS clave

Holoceno, palinomorfo, Provincia de Corrientes, sistema hidrográﬁco Iberá.

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

727

Bol. Soc. Argent. Bot. 57 (4) 2022

IntroductIon

relevant to construct modern analogs of this region

applied in palynological studies of surface and

The geological history of the region currently subsurface samples, although very few taxonomic

known as Iberá Wetlands in central-eastern works with illustration of palynomorphs published

South America, currently northeastern Argentina, up to today (Anzótegui & Garralla, 1985). Cuadrado

begins to have its own identity from the tectonic & Neiﬀ (1993) contributed with the ﬁrst analysis

movements that determined the rise of the Andean of disperse pollen from samples collected in

mountain range (Ramos, 1999). Orogeny unleashed dammed vegetation (“embalsados”) located in

compressive forces from the East that fractured the eastern region of Iberá Wetlands. These authors

powerful basaltic lava ﬂows belonging to the Solari- established that only upper levels yielded pollen

Serra Geral Formation (Cretaceous) accumulated grains representative of the current vegetation.

in the study region (Herbst & Santa Cruz, 1985). They also found that pollen grains are poorly

Thus, large blocks of rock generated and separated preserved, even physic and chemical features (e.g.

by several fault systems with dominant NE-SW low oxygen, low light, acid pH and high amounts of

and NW-SE strikes. The subsequent epirogenic organic matter) could have favored the preservation

movements gave rise to the independent adjustment of organic matter. Later, Garralla (1998) interpreted

of the megablocks, modeling the topography of that the region was vegetated by xero-halophyte

the subsoil covered by sedimentary ﬁll (Gentili forests at around 3,500 years BP.

&

Rimoldi, 1979). This generated in northeastern

Argentina, province of Corrientes, the extensive Characteristics of the study area

depression that crosses it almost entirely in a

Iberá Wetlands is one of the most important

NE-SW direction, on which important fluvial tropical wetlands of the biosphere, in terms of its

runoﬀ routes were developed. At the end of the extension and diversity of both animals and plants.

Pliocene, water excesses were concentrated in these The toponymy referred to “waters that shine” (Y verá

depressed lands, initiating its geological activity in in Guarani language). It comprises a complex water

the current Paraná River. Later tectonic movements system composed of, marshes, shallow lakes and

accentuated the vertical displacement of the subsoil interconnected river courses (Orfeo & Neiﬀ, 2008).

blocks, triggering changes in the runoﬀ design of

The lagoons correspond to extensive and shallow

surface waters, and being a consequence in the plains fed mainly by the rains (1,200 to 1,500

province of Corrientes, the Paraná River migrated mm year) and they are one of the most important

from South to North until it occupied its current wetlands in Latin America due to its extension with

2

position, which evidences a clear structural control more than 12,000 km . It is located in the central part

(

Iriondo, 1994). The central depression of the of Corrientes with a major axis NE-SW direction

Corrientes territory disconnected from the Paraná (Fig. 1), where a complex association of lentic and

River, in the surrounding of the town of Ituzaingó, lotic environments are vaguely and transitionally

where the old river valley transformed into a delineated (Neiff, 1997). Its extension into the

shallow basin, was ineﬃcient to evacuate excess Paraguay Republic “Esteros de Ñeembucú”, bearing

2

of water. This gave rise to the development of a about 45,000 km , are developed in its western area

complex association of lentic environments fed and conﬁrms the great inﬂuence of the rivers in the

fundamentally by precipitation and probably by maintainance of its ﬂuvial plains such as the large

the Paraná River due to subterranean transﬂuence Paraná (Neiﬀ, 2004).

(Orfeo, 2005). The plant communities of the study area in the

Therefore a complex association of lentic and Iberá Wetlands are included in the eastern district

lotic environments distributed over large areas of of the Chaco province. Its humid sandy plains are

the Iberá Wetlands in eastern Corrientes developed composed of typical savanna of Andropogon lateralis

during the Quaternary (Neiﬀ, 1997; Iriondo, 2010), Nees (Poaceae), accompanied by Cyperaceae and

from which taxonomic works of modern pollen other species of Poaceae, developed in more or less

grains of ﬂoristic biomes carried out mostly in elevated sectors well-drained; in depressions with

the last decades (e.g. Fernandez Pacella, 2013, slow permeability, this vegetation is sometimes

2

014). This botanical and ecological information is transformed into marshes (Carnevalli, 1994). Deeper

728

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

Fig. 1. Map of the western margin of the Iberá Wetland showing the sequence of the lakes studied, and cal

BP age range for each lake (from Fernandez Pacella & Lara, 2019).

sand areas (“espartillares”), that are well-drained (Fernandez Pacella et al., 2011, 2013; Fernandez

soils of pluvial in origin, are colonized by Elionurus Pacella & Lara, 2019) which represented mostly

muticus (Spreng.) Kuntze (Poaceae), as well as marsh-herbaceous vegetation under humid

grasslands of Sorghastrum agrostoides (Speg.) condition between 5968-5000 years cal BP. In

Hitchc., Paspalum sp. and Spartina sp. (Poaceae) agreement with Iriondo (1994), drier conditions

generally located in ﬂoodable soils like streams and were determined from 5000 to 3506 years cal BP

marshes. The hygrophilous forest (e.g. Poaceae, with a decrease in the species frequency typical of

Leguminosae) extended over wavy red sands wet environments and increase of xero-halophyte

(Carnevalli, 1994).

vegetation. This author pointed out that the

inﬂuence of template climate in Patagonia would

Paleoenvironment during the Holocene in the study have aﬀected the southern region of Corrientes

area evidencing changes in landscapes. The clogging of

Palynomorphs were identified in lacustrine the water body occurred from 3484 years cal BP

sediments of northwestern of the Iberá Wetlands onwards when herbaceous vegetation dominated.

729

Bol. Soc. Argent. Bot. 57 (4) 2022

The appearance of arboreal pollen indicates the

Thematerialprocessedandhousedinthelaboratory

beginning of the modern hygrophilous forest. at the CECOAL (Centro de Ecología Aplicada del

Those paleoenvironmental changes of alternating Litoral) of CONICET-UNNE (Universidad Nacional

wetter and drier episodes recorded in this region del Nordeste), in the collection “Dr. Rafael Herbst”.

are supported by ﬁelds of eolian dunes in Iberá

Wetlands, developed during the drier periods in the 30-50 pollen grains for each taxon, and it was done

late Holocene (Iriondo, 2010). under optical microscope NIKON ECLIPSE E100

The morphological description was based on

The objective of this contribution is to illustrated with a camera Nikon 590CU (40x and

provide the ﬁrst morphological documentation of 100x magniﬁcation). We followed the classiﬁcation

angiosperms, ferns, lycophytes and bryophytes of Curtis et al. (2001) and Raven et al. (1991) for

from core sediments of six lakes of the Holocene pollen grains, and genera included in the APG VI

western margin of the Iberá Wetlands. Species from Classiﬁcation System (2017) and PPG I (2016).

local versus extra-local vegetation diﬀerentiated in Terminology used for descriptions of pollen grains

the taxonomic description in order to contribute to and spores is based on Kremp (1965), Nilson &

a better understanding of the ﬂoral composition of Praglowski (1992), Punt et al. (2007), and Sáenz

the Iberá Wetlands and environmental changes yet Laín (2004). The reference collection PAL-CTES

documented during the Holocene.

of the UNNE and specialized literature were used

to identify pollen grains and spores (Acevedo &

Anzótegui, 1998; Anzótegui & Ferrucci, 1998;

Anzótegui, 2001; Anzótegui & Caccavari, 2001;

Anzótegui & Mautino, 2001a, b; Bhattacharya et al.,

materIalS and methodS

The taxonomic analysis of pollen grains and 2009; Caccavari & Dome, 2001, 2006; Cuadrado,

spores presented in this contribution is a new 1998a, b, c; Fernandez Pacella & Canteros, 2014;

study derived from previous studies carried out by Fernandez Pacella et al., 2014a, b; Fuertes &

Fernandez Pacella et al. (2011), Fernandez Pacella Rodríguez, 2009; Garralla & Cuadrado, 2001;

(2013) and Fernandez Pacella & Lara (2019) in Markgraf & D`Antoni, 1978; Pire et al., 2006).

which paleoenvironmental interpretations based on Local taxa refer to those species that are part

main palynologic groups addressed. These works of the natural vegetation of the Iberá Wetlands

included the study of core samples obtained with (Arbo & Tressens, 2002) whereas the extra-local

a Livingston-type sampler from six lakes in their taxa are those that do not belong to the Iberá. The

central and deepest parts on the western margin of latter species found in low percentages in the Tilia

the Iberá Wetlands. The interval studied of each diagrams of the lakes analyzed by Fernandez Pacella

1

4

core dated by C spans c. 6000 to 3000 Years BP et al. (2011) and Fernandez Pacella & Lara (2019).

Fig. 1). Samples were taken each 5 cm from ﬁve They achieved to depocenters by means of wind

(

lakes (San Juan, 75 cm, 27º 40´ 0´´ S 57º 11´ 53´´ currents coming from farther regional vegetation

W; San José, 85 cm, 27º 45´ 47´´ S 57º 13´ 20´´ W; of N, NW and W of Corrientes and E of Chaco

Lake “C”, 110 cm, 28º 10´ 41´´ S 57º 55´ 42´´ W; (Carnevali, 1994; Fontana, 2018). This is due to the

Rodeo Porá, 120 cm, 28° 30´ 30´´ S 58° 2´ 3´´ W; Iberá Wetlands was probably a closed basin made up

San Ignacio, 145 cm, 28° 32´ 32´´ S 58° 28´ 27´´ of an extensive mosaic of lentic bodies fed by rain

W). From the San Sebastián lake (24 cm, 27º 52´ and without connection with rivers or streams during

5

8´´ S 57º 20´ 34´´ W) the interval of sampling used the Mid-Late Holocene (Neiﬀ, 2004; Orfeo, 2005).

was each 2 cm.

The palynological maceration followed usual

techniques of Faegri & Iversen (1989), which reSultS

consisted on deﬂocculation of clays with (NaPO )

3

6

1

0%, elimination of humic acids with NaOH 5%,

Fifty-ﬁve palynomorphs were recorded in the

elimination of carbonates with HCl 10%, separation study area of the Iberá Wetlands. Forty-six of them

of organic matter from inorganic material with are pollen types corresponding to 27 families of

heavy liquids (ZnCl ) and elimination of silicates angiosperms and nine trilete spore-types of ferns,

2

with HF.

lycophytes, and bryophytes that are described and

730

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

illustrated (Figs. 2-5). In the taxonomic description, Amaryllidaceae (Liliopsida, Asparagales)

helpful information to diﬀerentiate local and extra- Crinum americanum L., Sp. Pl. 1: 292. 1753. Fig.

local taxa for each site studied is included (Table 1).

The 46 recorded species with pollen are listed

alphabetically by family:

2G.

Morphology: Monosulcate pollen grain, radial

symmetry. Small size, suboblate, (PA) 12-15 µm,

EA) 16-18 µm. Heteropolar. Sulcus 2 x 4 µm. Exine

2 µm thick, sexine tectate, microechinate of ca. 1 μm

(

Acanthaceae (Magnoliopsida, Lamiales)

Justicia brasiliana Roth, Nov. Pl. Sp.: 17. 1821. (Medeanic et al., 2008).

Fig. 2A-C. Ecological procedence: herbaceous local species

Morphology: Diporate pollen grain, radial well represented in marshes of the natural vegetation

symmetry. Circular amb, (PA) 17-84 µm, prolate of the Iberá Wetlands (Boelcke, 1992; Arbo &

EA) 14-43 µm. Lalongate endoapertures, 2-7 µm Tressens, 2002).

diam. Exine 1-5 µm thick, thinner in apertural area,

.5-1 μm thick, sexine semitectate, reticulated, Anacardiaceae (Magnoliopsida, Sapindales)

(

0

heterobrocate reticulum, with straight or sinuous Myracrodruon balansae (Engl.) Santin, Revista

walls and circular or polygonal lumens. Surrounding Brasil. Bot. 14: 135. 1991. Fig. 2H, I.

the aperture diﬀerentiate and contain 2-6 rows of

-10 islands (Pire et al., 2006).

Morphology: Tricolporate pollen grain, radial

symmetry. Subtriangular amb, subprolate, (PA) 21-28

4

Ecological procedence: grass or shrub well µm, (EA) 16-24 µm. Isopolar. Colpi 2.5 μm wide, ribs

represented in forests of S. balansae and M. 1.4-2 μm thick, lalongate endoapertures 1.4-2.1 x 7-10

balansae, corresponding to extra-local taxa of μm. Exine 1.4 µm thick. Sexine semitectate. Surface

Corrientes (Fontana, 2018). Native species of striate, short and wide striations lesser than 2.5 μm in

Chacoan and Paranaense ﬂoristic provinces (Anton length, and larger than 1.7 μm in width, walls 0.5 μm

&

Zuloaga, 2021).

wide and 0.2-0.3 μm high (Anzótegui, 2001).

Ecological procedence: native extra-local tree

represented in forests of S. balansae and M. balansae,

Amaranthaceae (Magnoliopsida, Caryophyllales)

Amaranthus muricatus (Moq.) Hieron., Bol. of the natural vegetation of Corrientes (Boelcke,

Acad. Nac. Ci. 4: 421. 1882. Fig. 2D. 1992; Fontana, 2018) and other region of Chacoan

Morphology: Pantoporate pollen grain, radial and Paranaense ﬂoristic provinces (Anton & Zuloaga,

symmetry, spheroidal, small size, diameter 19- 2021).

2

2 µm. Apolar. Circular pores, 28-45, with thick

margin, distributed all over the grain surface, Schinopsis balansae Engl., Bot. Jahrb. Syst. 6: 286.

-2 μm in diameter. Exine 1.2 a 1.5 µm thick. 1885. Fig. 2J.

Microreticulate (Cuadrado, 1998a). Morphology: Tricolporate pollen grain, radial

1

Ecological procedence: herbaceous species symmetry. Prolate, (PA) 32-35 µm, (EA) 24-27 µm.

frequent in natural grasslands of the Iberá Wetlands Isopolar. Colpi 1-2.5 μm maximum equatorial in

(

Arbo & Tressens, 2002) and mainly in North- width, acute or blunt apices, ribs 1.4-3.5 μm thick at

central Argentina (Anton & Zuloaga, 2021).

the equator. Lalongate endoapertures 3-4 x 7-8 μm.

Exine 0.7-2 µm thick. Surface striate, long striations,

Gomphrena perennis L., Sp. Pl. 1: 224. 1753. Fig. longer than 2.5 μm and 1 μm or less in width, walls

2

E, F.

Morphology: Pantoporate pollen grain, radial

slightly taller than 0.5 μm in height (Anzótegui, 2001).

Ecological procedence: native extra-local species

symmetry. Spheroidal, 14-22 µm. Apolar. Pores of the natural vegetation of Corrientes in forests of S.

0-37, 1.2-2.3 μm diam. Exine 1.5-3.5 µm thick, balansae and M. balansae (Fontana, 2018) belonging

metareticulate, with hexagonal elements (Cuadrado, to the Chacoan ﬂoristic province (Anton & Zuloaga,

998a). 2021).

Ecological procedence: herbaceous species in

grasslands of the Iberá Wetlands (Arbo & Tressens, Schinus longifolia (Lindl.) Speg., Cat. Descr.

002), and North-central region of Argentina Maderas: 413. 1910. Fig. 2K.

Anton & Zuloaga, 2021). Morphology: Tricolporate pollen grain, radial

3

1

2

(

731

Bol. Soc. Argent. Bot. 57 (4) 2022

Fig. 2. Pollen of Justicia, Amaranthus, Gomphrena, Crinum, Myracrodruon, Schinopsis, Schinus, Syagrus,

Alnus, Tecoma, Celtis, Anthemis, Baccharis, Bidens, Conyza, and Gaillardia. A: Justicia brasiliana,

equatorial view in superior focus (100x). B: Equatorial view in optical section (100x). C: Equatorial view

in optical section (100x). D: Amaranthus muricatus, general view in superior focus where endoapertures

are appreciated (100x). E: Gomphrena perennis, general view in optical section (100x). F: General view

in superior focus (100x). G: Crinum americanum, polar view in optical section (100x). H: Myracrodruon

balansae, equatorial view in superior focus (100x). I: Equatorial view in optical section (100x). J: Schinopsis

balansae, equatorial view in optical section (100x). K: Schinus longifolia, equatorial view in optical section

(100x). L: Syagrus romanzoﬃana, polar view in optical section (100x). M: Alnus acuminata, polar view in

optical section (100x). N: Tecoma stans, polar view in optical section (100x). O: Celtis iguanaea, equatorial

view (100x). P: Anthemis cotula, polar view in superior focus (100x). Q: Baccharis trimera, polar view in

optical section (100x). R: Bidens subalternans, polar view (100x). S: Conyza pampeana, polar view in

superior focus (100x). T: Gaillardia megapotamica, equatorial view (100x). Scales= 10 µm.

732

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

Table 1. Pollen and spores procedence for each site studied. Abreviatures= L: local taxa; EL: extra-local

taxa.

San

Ignacio Porá

Rodeo Lake

San

José

San

Juan

Species or genus

Alnus acuminata

Procedence

“C” Sebastián

L

X

Amaranthus muricatus

Anadenanthera colubrina

Anthemis cotula

X

EL

L

X

Anthoceros aﬀ. lamellatus

Baccharis trimera

L

X

Bidens subalternans

Calliandra parvifolia

Celtis iguanaea

L

X

EL

L

X

Chrysophyllum marginatum

Cissus verticillata

L

X

L

X

Conyza pampeana

L

X

Crinum americanum

Cyathea aﬀ. atrovirens

Cypella herbertii

L

X

L

X

L

X

Cyperus rotundus

L

X

Echinochloa crus-galli

Echinochloa polystachya

Empetrum aﬀ. rubrum

Eryngium elegans

L

X

Gaillardia megapotamica

Gomphrena perennis

Gutierrezia resinosa

L

X

Hymenachne

pernambucensis

L

X

Imperata brasiliensis

Jamesonia ﬂexuosa

Justicia brasiliana

L

X

EL

L

X

Ludwigia peploides

Mikania cordifolia

X

L

Myracrodruon balansae

Myrcianthes pungens

Myriophyllum aquaticum

Pellaea sp.

EL

L

X

L

X

Phaeoceros bulbiculosus

Phaeoceros tenuis

L

X

L

Phlegmariurus aﬀ.

mandiocanus

L

X

Phyllostylon rhamnoides

Pisonia aculeata

EL

X

733

Bol. Soc. Argent. Bot. 57 (4) 2022

San

Ignacio Porá

Rodeo Lake

San

José

San

Juan

Species or genus

Plinia rivularis

Procedence

“C” Sebastián

L

X

Polygala leptocaulis

Polygonum acuminatum

Polygonum convolvulus

Prosopis alba

X

L

X

L

EL

L

X

Schinopsis balansae

Schinus longifolia

X

Selaginella aﬀ. marginata

Senecio bonariensis

Senegalia bonariensis

Serjania perulacea

Sphagnum sp.

X

L

X

EL

L

X

Syagrus romanzoﬃana

Tecoma stans

EL

L

X

Thinouia mucronata

Trichilia elegans

L

X

EL

L

X

Typha domingensis

X

symmetry. Prolate, (PA) 18-21 µm, (EA) 14-16 µm. Mainly endemic of Pampean and Paranaense ﬂoral

Isopolar. Colpi 1-2.5 μm wide, with 1.4-3.5 μm provinces (Anton & Zuloaga, 2021).

thick ribs at the equator limited by a small psilate

margin of 2 μm wide. Endoapertures 1.4 x 4 µm in Betulaceae (Magnoliopsida, Fagales)

size. Exine 0.7-2.5 µm thick. Surface striate, long Alnus acuminata Kunth, Nov. Gen. Sp. (quarto

striations, approximately more than 2.5 μm long ed.) 2: 20. 1817. Fig. 2M.

(Anzótegui, 2001).

Morphology: Pentazonoporate pollen grain of

Ecological procedence: extra-local shrub or radial symmetry. Medium size, pentagonal amb

tree well represented in the natural palm grove (EA) 40-43 µm. Isopolar. Pores 3-5 µm, protrudent

vegetation of Corrientes (Fontana, 2018) and in and vestibular. Exine 2 µm thick, psilate (Markgraf

Chacoan and Paranaense ﬂoral provinces (Anton & & D`Antoni, 1978).

Zuloaga, 2021).

Ecological procedence: extra-local tree

represented in forests of S. balansae and M.

balansae of Corrientes (Boelcke, 1992; Fontana,

Arecaceae (Liliopsida, Arecales)

Syagrus romanzoffiana (Cham.) Glassman, 2018).

Fieldiana, Bot. 31: 382. 1968. Fig. 2L.

Morphology: Monosulcate pollen grain, bilateral Bignoniaceae (Magnoliopsida, Lamiales)

symmetry. Medium size, oblate, (PA) 15-17 µm, Tecoma stans (L.) Juss. ex Kunth, Nov. Gen. Sp.

(

EA) 37-39 µm. Heteropolar. Sulcus 35.4 μm long. (quarto ed.) 3: 144. 1819. Fig. 2N.

The exine is 1.6 μm thick, sexine tectate and psilate Morphology: Brevicolpate pollen grain, radial

Trigo & Fernández, 1995; Bauermann et al., 2010). symmetry. Subtriangular amb, (EA) 48-52 µm.

Ecological procedence: extra-local arboreal Isopolar. Exine 3 µm thick, sexine semitectate.

species well represented in small palm stocks and Microreticulate (Markgraf & D`Antoni, 1978).

riparian natural vegetation in Corrientes (Boelcke, Ecological procedence: shrub or small tree

992; Carnevalli, 1994; Cabral & Castro, 2007). represented in the natural vegetation of the Iberá

(

1

734

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

Wetlands in hygrophilous forests (Fontana, 2018). 27-28 µm. Isopolar. Exine 2.5-3 µm thick, sexine

Native of North-central region of Argentina (Anton tectate, echinate, spines 7-7.5 µm (Markgraf &

&

Zuloaga, 2021).

D`Antoni, 1978).

Ecological procedence: herbaceous taxon

presents in grasslands of the Iberá Wetlands (Arbo

Celtidaceae (Magnoliopsida, Rosales)

Celtis iguanaea (Jacq.) Sarg., Silva 7: 64. 1895. & Tressens, 2002).

Fig. 2O.

Morphology: Triporate pollen grain, radial Conyza pampeana (Parodi) Cabrera, Man. Fl. Al.

symmetry. Subtriangular to circular amb, suboblate, Buenos Aires: 481. 1953. Fig. 2S.

(

PA) 9-30 µm, (EA) 11-32 µm. Isopolar. Pores

Morphology: Tricolporate pollen grain of radial

slightly sunken, circular 1-2 μm in diam., limited symmetry. Prolate to subcircular, (PA) 25-27 µm,

by a ring 0.5-2 μm thick. Exine 0.5-1.4 µm thick. (EA) 21-24 µm. Isopolar. Exine 4 µm thick. Sexine

Sexine tectate. Scabrate (Anzótegui & Mautino, tectate, echinate, spines 2 µm (Medeanic et al.,

2

001a).

Ecological procedence: shrub (creep) or tree

2008).

Ecological procedence: this genus is well

represented in hygrophilous forests of the Iberá represented in the natural vegetation of the Iberá

Wetlands and extra-local forests of S. balansae and Wetlands in grasslands (Arbo & Tressens, 2002).

M. balansae in Corrientes (Arbo & Tressens, 2002;

Fontana, 2018). Native of central-northern region Gaillardia megapotamica (Spreng.) Baker, Fl.

of Argentina (Anton & Zuloaga, 2021).

Bras. 6: 276. 1884. Fig. 2T.

Morphology: Tricolporate pollen grains of radial

symmetry. Medium, suboblate, (PA) 27-30 µm,

(EA) 34-37 µm. Isopolar. Exine 2 µm thick, sexine

Compositae (Magnoliopsida, Asterales)

Anthemis cotula L., Sp. Pl. 2: 894. 1753. Fig. 2P.

Morphology: Tricolporate pollen grains of radial tectate, echinate, with spines 4 µm long (Markgraf

symmetry. Spheroidal, (PA) 23-24 µm, (EA) 22-23 & D`Antoni, 1978).

µm. Isopolar. Exine 4.5 µm thick. Sexine tectate,

echinate, spines 4 to 5 μm (Alonso, 2014).

Ecological procedence: perennial grass or

subshrub well represented in grasslands of the Iberá

Ecological procedence: herbaceous local taxon Wetlands (Arbo & Tressens, 2002).

in grasslands of the Iberá Wetlands (Boelcke, 1992;

Arbo & Tressens, 2002) in Corrientes and widely Gutierrezia resinosa (Hook. & Arn.) S. F. Blake,

present in southern South America (Anton & Contr. U.S. Natl. Herb. 26: 232. 1930. Fig. 3A.

Zuloaga, 2021).

Morphology: Tricolporate pollen grain, radial

symmetry. Small size, spheroidal, (PA) 14-18 µm,

Baccharis trimera (Less.) DC., Prodr. 5: 425. (EA) 14-18 µm. Isopolar. Exine 2 µm thick. Sexine

836. Fig. 2Q. tectate, echinate, spines 1.5 µm long (Markgraf &

Morphology: Tricolporate, radial symmetry. D`Antoni, 1978).

Spheroidal, (PA) 15-16 µm, (EA) 14-15 µm. Ecological procedence: Gutierrezia Lag. is

1

Isopolar. Exine 4 µm thick. Sexine tectate, echinate, present in natural grasslands of the Iberá Wetlands

spines 2-2.5 µm long (Markgraf & D`Antoni, (Arbo & Tressens, 2002).

1

978).

Ecological procedence: this genus is well Mikania cordifolia (L. f.) Willd., Sp. Pl. 3: 1746.

represented in natural grassland vegetation of 1803. Fig. 3B.

the Iberá Wetlands (Arbo & Tressens, 2002) and

Morphology: Tricolporate pollen grains of radial

in North-central region of Argentina (Anton & symmetry. Medium size, subprolate (PA) 27-30 µm,

Zuloaga, 2021).

(EA) 22-25 µm. Isopolar. Exine 1 µm thick, sexine

tectate, echinate, spines 2 µm (Ybert et al., 2016).

Ecological procedence: perennial creeper well

represented in hygrophilous forest vegetation of

Bidens subalternans DC., Prodr. 5: 600. 1836.

Fig. 2R.

Morphology: Tricolporate pollen grain, radial Iberá Wetlands (Arbo & Tressens, 2002), as well

symmetry. Oblate-spheroidal, (PA) 26-28 µm, (EA) as in forests, savannas, riparian banks of rivers

735

Bol. Soc. Argent. Bot. 57 (4) 2022

and streams of South America (Anton & Zuloaga, Iberá Wetlands (Arbo & Tressens, 2002) also

2

021).

present in central-northern region of Argentina

Anton & Zuloaga, 2021).

(

Senecio bonariensis Hook. &Arn., J. Bot. (Hooker)

: 340. 1841. Fig. 3C.

Morphology: Tricolporate pollen grain, radial Cypella herbertii Herb., Bot. Mag. 53: sub t. 2637.

symmetry. Circular in (PA) 28-40 µm, prolate in 1826. Fig. 3H.

EA) 16-35 µm. Isopolar. Exine 2-5 µm thick. Morphology: Monosulcate pollen grain and

Sexine tectate, echinate, spines 2-3 µm (Markgraf bilateral symmetry. Heteropolar, suboblate, (PA)

D`Antoni, 1978). 40-54 µm, (EA) 45-61 µm. Sulcus 35-54 µm

Ecological procedence: Senecio L. is well long. Exine 1-2.2 μm thick, sexine semitectate and

represented in the natural vegetation of the Iberá microreticulate (Salgado, 2006).

3

Iridaceae (Liliopsida, Asparagales)

(

&

Wetlands in marsh and grassland, corresponding

Ecologic procedence: bulbous herbaceous local

to local taxa (Arbo & Tressens, 2002) and present species in marsh areas (Boelcke, 1992; Arbo

in Chacoan, Paranaense and Pampean floristic & Tressens, 2002). Endemic of Pampean and

provinces (Anton & Zuloaga, 2021).

Paranaense ﬂoral provinces (Anton & Zuloaga,

021).

2

Cyperaceae (Liliopsida, Poales)

Cyperus rotundus L., Sp. Pl. 1: 45. 1753. Fig. 3D. Leguminosae, Mimosoideae (Magnoliopsida,

Morphology: Monoporate pollen grain and radial Fabales)

symmetry. Heteropolar, prolate, (PA) 27-37 µm, Calliandra parvifolia (Hook. & Arn.) Speg.,

(EA) 16-24 µm. Exine 1.5 µm thick, ornamentation Revista Argent. Bot. 1: 193. 1926. Fig. 3I.

granular (Fernandez, 1987). Morphology: Polyads 185 x 112 μm diam.

Ecological procedence: perennial marsh grass usually of 8 pollen grains. Exine 2 µm thick,

well represented in the natural vegetation of the rugulate (Markgraf & D`Antoni, 1978).

Iberá Wetlands (Arbo & Tressens, 2002).

Ecological procedence: extra-local shrub

represented in the natural vegetation of Corrientes

province in forests of S. balansae and M. balansae

Ericaceae (Magnoliopsida, Ericales)

Empetrum aﬀ. rubrum Vahl ex Willd., Sp. Pl. 4: (Fontana, 2018).

7

13. 1806. Fig. 3E, F.

Morphology: Tetrad with tricolporate pollen Anadenanthera colubrina (Vell.) Brenan, Kew

grain, radial symmetry. Small, circular-subcircular, Bull. 10: 182. 1955. Fig. 3J.

(

EA) 19-22 µm. Isopolar. Exine 2 µm thick

Markgraf & D`Antoni, 1978).

Morphology: Polyads spheroidal to ellipsoidal

of 12 pollen grains irregularly arranged, or of 16

Ecological procedence: subshrub represented pollen grains regularly arranged. Exine 1.3 µm

in marshes of the natural vegetation of the Iberá thick. Verrucate, 1 μm diam. warts (Caccavari &

Wetlands (Carnevalli, 1994; Arbo & Tressens, Dome, 2006).

2

002).

Ecological procedence: extra-local tree

represented in forests of S. balansae and M.

balansae of Corrientes province (Fontana, 2018).

Haloragaceae (Magnoliopsida, Saxifragales)

Myriophyllum aquaticum (Vell.) Verdc., Kew

Bull. 28: 36. 1973. Fig. 3G.

Prosopis alba Griseb., Abh. Königl. Ges. Wiss.

Morphology: Tetrazonoporate pollen grain with Göttingen 19: 131. 1874. Fig. 3K.

radial symmetry. Subcircular-angular amb, oblate

Morphology: Tricolporate pollen grain of radial

to spheroidal (PA) 29-38 µm, (EA) 34-41 µm. symmetry. Small to medium size, prolate, (PA)

Pores 4-5 μm diam., protrudent, with thickening of 22-36 µm, (EA) 18.5-33 µm. Isopolar. Long colpi

the endexine. Exine 1.5 µm thick. Sexine tectate, 17-30 µm and pores 4-6 μm diam. Exine 1-2 µm

scabrate (Díez Dapena, 1988).

Ecological procedence: perennial aquatic herb al., 2014b).

well represented in the natural vegetation of the Ecological procedence: extra-local tree

thick, sexine tectate, scabrate (Fernandez Pacella et

736

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

Fig. 3. Pollen of Gutierrezia, Mikania, Senecio, Cyperus, Empetrum, Myriophyllum, Cypella, Calliandra,

Anadenanthera, Prosopis, Senegalia, Trichilia, Myrcianthes, and Plinia. A: Gutierrezia resinosa, equatorial

view in optical section (100x). B: Mikania cordifolia, polar view in optical section (100x). C: Senecio

bonariensis, polar view in optical section (100x). D: Cyperus rotundus, equatorial view in optical section

(

100x). E: Empetrum aﬀ. Rubrum, poliade in optical section (100x). F: Poliade in superior focus (100x). G:

Myriophyllum aquaticum, polar view in superior focus (100x). H: Cypella herbertii, polar view in optical section

100x). I: Calliandra parvifolia, poliade (100x). J: Anadenanthera colubrina, poliade (100x). K: Prosopis alba,

(

polar view in superior focus (100x). L: Senegalia bonariensis, poliade (100x). M: Trichilia elegans, equatorial

view in optical section (100x). N: Equatorial view in superior focus (100x). O: Myrcianthes pungens, polar

view in superior focus (100x). P: Plinia rivularis, polar view in superior focus (100x). Scales= 10 µm.

represented in forests of S. balansae and M.

Morphology: Polyads of 16 pollen grains with

balansae of Corrientes (Boelcke, 1992; Fontana, regular arrangement, major axis (34-42 μm), minor

2018).

axis (22-30 μm). Exine 1.55-2.55 µm thick. Sexine

tectate. Infratectum with columellae 0.5-0.9 μm

Senegalia bonariensis (Gillies ex Hook. & Arn.) long. Ornamentation slightly granulate (Caccavari

Seigler & Ebinger, Phytologia 88: 50. 2006. Fig. 3L. & Dome, 2001).

737

Bol. Soc. Argent. Bot. 57 (4) 2022

Ecological procedence: tree or shrub present Exclusive of Entre Ríos, Corrientes and Misiones

in hygrophilous forests of the Iberá Wetlands belonging to the Paranaense ﬂoristic province

(Boelcke, 1992; Fontana, 2018).

(Anton & Zuloaga, 2021).

Meliaceae (Magnoliopsida, Sapindales)

Nyctaginaceae (Magnoliopsida, Caryophyllales)

Trichilia elegans A. Juss., Fl. Bras. Merid. Pisonia aculeata L., Sp. Pl. 2: 1026. 1753. Fig.

(

quarto ed.) 2: 79, t. 98. 1829. Fig. 3M, N.

Morphology: Tetracolporate pollen grain,

4A, B.

Morphology: Tricolporate pollen grain of

radial symmetry. Small size, quadrangular radial symmetry. Medium size, prolate, (PA) 34-

amb (PA) 17-25 µm, subprolate (EA) 14-21 37 µm, (EA) 24-27 µm. Isopolar. Colpi 32-35 μm

µm. Isopolar. Colpi 15-22 μm long, lalongate long. Pores 2-3 μm diam. Exine 1-1.5 µm thick,

endoapertures with ring thickening 1.4-3 μm sexine semitectate, microreticulate (Oliveira &

thick. Exine 1.4 μm thick and microreticulate Santos, 2014).

ornamentation (Oliveira & Santos, 2014).

Ecological procedence: shrub or creeper

Ecological procedence: extra-local tree species present in natural forests of Schinopsis

represented in forests of S. balansae and M. balansae Engl. and Myracrodruon balansae

balansae of the natural vegetation of Corrientes (Engl.) Santin in Corrientes province (extra-local

(

&

Fontana, 2018) and noreastern Argentina (Anton taxon) (Carnevalli, 1994; Fontana, 2018), and

Zuloaga, 2021). northern region of Argentina (Anton & Zuloaga,

021).

2

Myrtaceae (Magnoliopsida, Myrtales)

Myrcianthes pungens (O. Berg) D. Legrand, Onagraceae (Magnoliopsida, Myrtales)

Bol. Fac. Agron. Univ. Montevideo 101: 52. Ludwigia peploides (Kunth) P.H. Raven,

1

968. Fig. 3O.

Reinwardtia 6: 393 1963[1964]. Fig. 4C.

Morphology: Parasintricolporate pollen

Morphology: Tricolporate pollen grain,

grain, radial symmetry. Small size, triangular, radial symmetry. Large size, subtriangular amb,

oblate, (PA) 9-17 µm, (EA) 12-21 µm. Isopolar. subprolate, (PA) 94-108 µm, (EA) 74-86 µm.

Narrow or wide colpi an apocolpial ﬁeld or Isopolar. Colpi 18 x 4 µm, endoapertures 20 x 16

island, of triangular form. Apocolpial ﬁelds with µm protrudent-vestibuled. Exine 3-4 µm thick.

edges and are of diﬀerent sizes in both poles. The surface of the ridges is rugulate-striated. The

Endoapertures 1.4-2 μm. Exine 0.7-3 µm thick, area between the ridges has an uneven coarse

sexine semitectate, microreticulate (Acevedo & rugulate appearance, due to the presence of

Anzótegui, 1998).

irregular striae (Cecotti Álvarez et al., 2017).

Ecological procedence: the natural vegetation

Ecological procedence: aquatic herbaceous

of Corrientes province in forests of S. balansae taxon well represented in fresh water bodies

and M. balansae (Boelcke, 1992; Fontana, 2018). of the Iberá Wetlands (Boelcke, 1992; Arbo

Native of North-central region of Argentina & Tressens, 2002). Native of central-northern

(

Anton & Zuloaga, 2021). region of Argentina (Anton & Zuloaga, 2021).

Plinia rivularis (Cambess.) Rotman, Bol. Soc. Poaceae (Liliopsida, Poales)

Argent. Bot. 24: 195. 1985. Fig. 3P.

Echinochloa crus-galli (L.) P. Beauv., Ess.

Morphology: Parasyntricolporate pollen grain, Agrostogr. 1: 53, 161, 169, pl. 11, f. 2. 1812. Fig.

radial symmetry. Triangular amb, oblate, (PA) 4D.

1

3-17 µm, (EA) 19-28 µm. Isopolar. Linear colpi,

Morphology: Monoporate pollen grain and

less than 1 μm wide, lalongate endoapertures 1 radial symmetry. Spheroidal amb, (PA) 46-49

x 2.3 µm. Exine 0.7-2 µm thick. Sexine tectate. µm, (EA) 46-49 µm. Heteropolar. Pore 2 μm

Psilate (Acevedo & Anzótegui, 1998).

in diameter, with annulus 1 µm thick. Exine

Ecological procedence: tree well represented 2 µm thick. Sexine tectate, psilate to scabrate

in hygrophilous forests of the Iberá Wetlands (Fernandez Pacella & Canteros, 2014).

(

Boelcke, 1992; Arbo & Tressens, 2002).

Ecological procedence: herbaceous species in

738

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

marsh and grasslands of natural vegetation of the Amer. 1: 130. 1838. Fig. 4H.

Iberá Wetlands (Boelcke, 1992; Arbo & Tressens, Morphology: Pantozonocolporate pollen grain

002) and widely distributed in Argentina (Anton with radial symmetry. Subcircular amb (PA)

2

&

Zuloaga, 2021).

20-30 µm, subprolate amb (EA) 18-21 µm.

Isopolar. Colpi 8-15, 17-25 μm long. Lalongate

Echinochloa polystachya (Kunth) Hitchc., endoaperture 2 x 4 µm. Exine 0.7-2.5 µm thick.

Contr. U.S. Natl. Herb. 22: 135. 1920. Fig. 4E.

Sexine tectate. Psilate (Cuadrado, 1998c).

Morphology: Monoporate pollen grain and

Ecological procedence: herbaceous species

radial symmetry. Spheroidal amb, (PA) 53-57 well represented in marshes and grasslands of the

µm, (EA) 60-62 µm. Heteropolar. Pore with Iberá Wetlands (Boelcke, 1992; Arbo & Tressens,

annulus 2 µm in diameter. Exine 2 µm thick, 2002).

sexine tectate, psilate to scabrate (Fernandez

Pacella & Canteros, 2014).

Polygonaceae (Magnoliopsida, Caryophyllales)

Ecological procedence: herbaceous taxon, Polygonum acuminatum Kunth, Nov. Gen. Sp.

local of marsh and grasslands in the natural (quarto ed.) 2: 178. 1818. Fig. 4I, J.

vegetation of the Iberá Wetlands (Arbo &

Morphology: Triporate pollen grain with radial

Tressens, 2002). Extra-local species present symmetry. Circular amb, (EA) 59-62 µm. Exine 6

in Paranense, Chacoan and Pampean floral µm thick. Sexine 1.5-2 µm thick, reticulate, walls

provinces of Argentina (Anton & Zuloaga, 2021). 1.5 μm thick, simplicolumellate, lumen 10 μm

wide (Ybert et al., 2018).

Hymenachne pernambucensis (Spreng.)

Zuloaga, Amer. J. Bot. 90: 817. 2003. Fig. 4F.

Ecological procedence: herbaceous species

well represented in hygrophilous forests of the

Morphology: Monoporate pollen grain, radial Iberá Wetlands (Fontana, 2018).

symmetry. Prolate-spheroidal, (PA) 42-44 µm,

(

EA) 36-39 µm. Heteropolar. Pore 3 µm in Polygonum convolvulus L., Sp. Pl. 1: 364. 1753.

diameter. Exine 2 µm thick. Sexine tectate, psilate Fig. 4K.

to scabrate (Fernandez Pacella & Canteros, Morphology: Triporate pollen grain with

014). radial symmetry. Circular-subcircular amb, (EA)

Ecological procedence: H. pernambucensis 39-42 µm. Isopolar. Exine 3 µm thick, reticulate,

2

local genus is well represented in marsh and walls 1 μm thick, simplicolumellate, irregular

grasslands (Arbo & Tressens, 2002). Present lumen 2 x 2 μm wide (Ybert et al., 2018).

in Paranense and Chacoan ﬂoral provinces of

Argentina (Anton & Zuloaga, 2021).

Ecological procedence: herbaceous species

represented in hygrophilous forests of the Iberá

Wetlands (Fontana, 2018).

Imperata brasiliensis Trin., Mém. Acad. Imp.

Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2: 331. Sapindaceae (Magnoliopsida, Sapindales)

832. Fig. 4G. Serjania perulacea Radlk., Consp. Sect. Sp.

Morphology: Monoporate pollen grain, radial Serjan.: 11. 1874. Fig. 4L, M.

symmetry. Spheroidal amb, (PA) 21-37 µm, (EA) Morphology: Hemisyntricolporate pollen

8-33 µm. Heteropolar. Pore 1-4 μm diameter grain, radial symmetry. Subtriangular amb,

1

with a ring 1 to 6 μm thick. Exine 1-3 μm thick, oblate, (PA) 8-40 µm, (EA) 6-58 µm. Heteropolar.

tectate, psilate to scabrate (Fernandez Pacella & Colpi 1-2 μm wide. Exine 0.7-2.5 µm thick,

Canteros, 2014).

sexine semitectate, microreticulate (Anzótegui &

Ecological procedence: herbaceous species Ferrucci, 1998).

well represented in marsh and grassland areas

Ecological procedence: extra-local perennial

of the natural vegetation of the Iberá Wetlands creeper represented in the forests of Schinopsis

(

Boelcke, 1992; Arbo & Tressens, 2002).

balansae and Myracrodruon balansae, natural

vegetation of Corrientes (Fontana, 2018) and

other provinces of northern Argentina: Chaco,

Polygalaceae (Magnoliopsida, Fabales)

Polygala leptocaulis Torr. & A. Gray, Fl. N. Formosa, Jujuy, Salta (Anton & Zuloaga, 2021).

739

Bol. Soc. Argent. Bot. 57 (4) 2022

Fig. 4. Pollen of Pisonia, Ludwigia, Echinochloa, Hymenachne, Imperata, Polygala, Polygonum, Serjania,

Thinouia, Chrysophyllum, and Typha. A: Pisonia aculeate, equatorial view in optical section (100x). B:

Equatorial view in superior focus (100x). C: Ludwigia peploides, polar view in optical section (100x). D:

Echinochloa crus-galli, equatorial view in superior focus (100x). E: Echinochloa polystachya, equatorial

view in superior focus (100x). F: Hymenachne pernambucensis, equatorial view in optical section (100x).

G: Imperata brasiliensis, equatorial view in superior focus (100x). H: Polygala leptocaulis, equatorial view in

superior focus (100x). I: Polygonum acuminatum, polar view in superior focus (100x). J: Polar view in optical

section (100x). K: Polygonum convolvulus, polar view in superior focus (100x). L: Serjania perulacea, polar

view in inferior focus (100x). M: Polar view in superior focus (100x). N: Thinouia mucronata, polar view in

superior focus (100x). O: Chrysophyllum marginatum, equatorial view in optical section (100x). P: Typha

domingensis, equatorial view in superior focus (100x). Scales= 10 µm.

Thinouia mucronata Radlk., Sitzungsber. Math.- to substraight. Isopolar. Colpi (1.4-4 μm wide).

Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: Lalongate endoapertures 1.4 x 4.3 μm. Exine 1.4-

2

82. 1878. Fig. 4N. 2.1 thick. Psilate to slightly scabrate (Anzótegui &

Morphology: Tricolporate pollen grain, radial Ferrucci, 1998).

symmetry. Subtriangular amb, oblate, (PA) 11- Ecological procedence: local perennial creeper

7 µm, (EA) 23-27 µm. Sides slightly convex well represented in hygrophilous forest vegetation

2

740

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

of the Iberá Wetlands (Arbo & Tressens, 2002; Fontana, 2018). Native of Chacoan and Yungas

Boelcke, 1992) and in northern Argentina (Anton & ﬂoristic provinces of Argentina (Anton & Zuloaga,

Zuloaga, 2021).

2021).

Sapotaceae (Magnoliopsida, Ericales)

Umbeliferae (Magnoliopsida, Apiales)

Chrysophyllum marginatum (Hook. & Arn.) Eryngium elegans Cham. & Schltdl., Linnaea 1:

Radlk., Act. Occ. Exp. Univ. Anvers Coin. Exp. Int. 348. 1826. Fig. 5C.

Hort.: 170. 1887. Fig. 4O.

Morphology: Tricolporate pollen grain, radial

Morphology: Tetracolporate pollen grain with symmetry. Prolate, (PA) 28-36 µm, (EA) 18-23

radial symmetry. Subtriangular amb, prolate, (PA) µm. Isopolar. Narrow and long colpi, relatively

2

±

0-24 µm, (EA) 14-16 µm. Isopolar. Narrow colpi large pore. Exine 1 µm thick. Sexine semitectate,

14 μm long. Endoapertures 2 x 3 µm with an perforate, microreticulate (Alonso, 2014).

annular thickening of 2 μm given by protruding Ecological procedence: local herb in marshes

endexine. Exine 3 µm thick, columellate infratectal of the Iberá Wetlands (Boelcke, 1992; Arbo &

layer observed. At the poles, the exine is 2 μm, the Tressens, 2002), and elsewhere in Corrientes

columellae being longer than the thick of the tectum (Fontana, 2018) and North-central Argentina

and the nexine 1 μm; at the equator the sexine thins, (Anton & Zuloaga, 2021).

until approximately 1 μm and the nexine increases

its thickness to 2 μm. Surface scabrate (Cuadrado, Vitaceae (Magnoliopsida, Vitales)

1

998b).

Ecological procedence: shrub or small tree Taxon 33: 727. 1984. Fig. 5D-F.

well represented in forests of the Iberá Wetlands Morphology: Tricolporate pollen grain, radial

and Norht-central region of Argentina (Arbo & symmetry. Subtriangular amb, subprolate, (PA)

Cissus verticillata (L.) Nicolson & C.E. Jarvis,

Tressens, 2002; Anton & Zuluoga, 2021).

75-77 µm, (EA) 58-60 µm. Isopolar. Colpi long,

maximum width of 4.5 μm and ribs 2.8-4.5 μm

thick at the equator. Lalongate endoapertures 5-8

Typhaceae (Liliopsida, Poales)

Typha domingensis Pers., Syn. Pl. 2: 532. 1807. x 3-4 μm to circular 3 μm diam. Exine 2-4 µm

Fig. 4P.

Morphology: Monoporate pollen grain, radial

thick, reticulate (Anzótegui & Caccavari, 2001).

Ecological procedence: perennial creeper

symmetry. Spheroidal amb, (PA) 17-20 µm, (EA) represented in hygrophilous forest of the natural

7-20 µm. Heteropolar. Exine 2.5 μm thick. Sexine vegetation of the Iberá Wetlands (Fontana, 2018)

columellate, wall 0.25 μm thick, reticulate (Alonso, and in central-northern of Argentina (Anton &

1

2

014).

Ecological precedence: robust emergent aquatic

herbaceous local taxon well represented in marshes

Zuloaga, 2021).

The nine recorded species with spores are

of natural vegetation of the IberáWetlands (Boelcke, listed alphabetically by family:

992; Arbo & Tressens, 2002).

1

A n t h o c e ro t a c e a e ( A n t h o c e ro t o p s i d a ,

Anthocerotales)

Ulmaceae (Magnoliopsida, Rosales)

Phyllostylon rhamnoides (J. Poiss.) Taub., Oesterr. Anthoceros aﬀ. lamellatus Steph., Sp. Hepat.

Bot. Z. 11: 409. 1890. Fig. 5A, B.

(Stephani) 5: 1000. 1916. Fig. 5G, H.

Morphology: Tetra-pentazonoporate pollen

Morphology: Trilete spore. Polar view

grain, radial symmetry. Spheroidal, (PA) 36-39 µm, subtriangular. Laesura with straight rays reaching

EA) 36-39 µm. Isopolar. Circular pores 1-3 μm in the equator. Exospore 3.5 µm thick, smooth

(

diam., some of them arranged in a subequatorial inner proximal surface, bacullate distal surface

plane. Exine 1 µm thick. Psilate to verrucate and equatorial amb. Size: 48-76 µm (Gradstein,

(

Anzótegui & Mautino, 2001b).

Ecological procedence: extra-local tree well

2018).

Ecological procedence: cosmopolitan genus is

represented in forests of S. balansae and M. represented on compact rock surfaces exposed to

balansae of Corrientes province (Boelcke, 1992; light (Peñaloza-Bojacá et al., 2020).

741

Bol. Soc. Argent. Bot. 57 (4) 2022

Cyatheaceae (Polypodiopsida, Cyatheales)

the equator. Exospore 3.5 µm thick. Granular.

Cyathea aﬀ. atrovirens (Langsd. & Fisch.) Domin, Size: 48-76 µm (Prieto & Quattrocchio, 1993).

Rozpr. Kral. Ceske Spolecn. Nauk, Tr. Mat.-Prir. 2:

62. 1929. Fig. 5I.

Ecological procedence: cosmopolitan genus is

represented on compact, moist soil, and on rock in

2

Morphology: Trilete spore. Polar view moist environments, exposed or partially shaded

triangular. Laesura 35-40 µm long. Exospore 1.4 microhabitats (Peñaloza-Bojacá et al., 2020).

µm thick. Psilate or scabrate. Size: 43-53 µm

(

Contreras-Duarte et al., 2006).

Ecological procedence: tree to shrub and Jamesonia ﬂexuosa (Kunth) Christenh., Phytotaxa

herb on soils and rock substrate (rupicolous 19: 21. 2011. Fig. 5M.

Pteridaceae (Polypodiopsida, Polypodiales)

species) on roadsides, ravines, in abandoned

Morphology: Trilete spore. Polar view

ﬁelds, swamps and secondary forests of forests of subtriangular, with rounded angles. Sinuous

the Iberá Wetlands (Arbo & Tressens, 2002) and laesura, with small coalescent ridges, reaching

in Corrientes and Misiones provinces (Anton & to the inner margin of the cingulum. Exospore

Zuloaga, 2021).

including the cingulum is 5.5 µm thick. Size: 56-71

µm (Contreras-Duarte et al., 2006; Della & Prado,

2020).

Ecological procedence: this taxon is part of the

natural vegetation of the Iberá Wetlans (Arbo &

Lycopodiaceae (Lycopodiopsida, Lycopodiales)

Phlegmariurus aff. mandiocanus (Raddi) B.

Øllg., Rodriguésia 63: 480. 2012. Fig. 5J.

Morphology: Trilete spore. Polar view Tressens, 2002) and occurs in humid places inside

subtriangular, with rounded angles. Lesura with of cloud forest from southern Mexico to Bolivia,

straight rays reaching to the equator. Exospore 1.6 and north and southeast of Brazil to Uruguay

µm thick. Psilate on proximal face. Size: 28-38 (Della & Prado, 2020; Della et al., 2020).

µm. Illustrations of this taxon by Sersic (1983)

and comparison with other species of the genus Pellaea sp. Fig. 5N

consulted in Rincón et al. (2014).

Morphology: Trilete spore. Polar view

Ecological procedence: epiphyte present in triangular. Rays of the laesurae straight, 2/3 of

hygrophilous forests of Yungas and Paranaense radius to almost the internal margin of cingulum.

ﬂoral provinces in Argentina (Anton & Zuloaga, Exospore cingulum surrounds the spore, 5.4 μm

2

021; Windisch et al., 2015).

thick. Central part of distal face verrucate/rugulate,

low warts occasionally joined forming loins. Size:

N ot ot hy l adac e ae (A nt hoc e rot opsi da, 37-48 µm.

Notothyladales)

Comparisons: few specimens recorded herein

Phaeoceros bulbiculosus (Brot.) Prosk., Rapp. are similar to Pellaea cordifolia (Sessé & Moc.)

Comm., VIII Congr. Int. Bot.: 69. 1954. Fig. 5K.

A.R. Sm. (Arreguín-Sánchez et al., 1996; Pérez-

Morphology: Trilete spore. Polar view Jiménez et al., 2020) and Pellaea ovata (Desv.)

subtriangular. Laesura with straight rays reaching Weath. (Gómez et al., 2013) diﬀering in having

the equator. Exospore 2.3 µm thick. Granular/ smooth proximal and less ornamented distal face.

gemmate sculpture. Size: 32-38 µm. Illustrated

from Argentina (Morbelli et al., 2010).

Ecological procedence: this genus is including

terrestrial or rupicolous species, some of humid

Ecological procedence: cosmopolitan genus is shaded areas of hygrophilous forests and others

represented on compact, moist soil, and on rock in tolerate slight exposition to light in open areas.

moist environments, exposed or partially shaded Particularly, Pellaea cordifolia and P. ovata (Hirai

microhabitats (Peñaloza-Bojacá et al., 2020).

& Prado, 2021) are known in South America

Anton & Zuloaga, 2021; Arbo & Tressens, 2002).

(

Phaeoceros tenuis (Spruce) Hässel, Veröff.

Geobot. Inst. ETH Stiftung Rübel Zürich 91: 303. Selaginellaceae (Lycopodiopsida, Selaginellales)

986. Fig. 5L. Selaginella aﬀ. marginata (Humb. & Bonpl. ex

Morphology: Trilete spore. Polar view Willd.) Spring, Flora 21: 194. 1838. Fig. 5O.

subtriangular. Laesura with straight rays reaching Morphology: Trilete microspore. Polar view

1

742

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

Fig. 5. Palynomorphs of Phyllostylon, Eryngium, Cissus, Anthoceros, Cyathea, Phlegmariurus, Phaeoceros,

Jamesonia, Pellaea, and Sellaginella. A: Phyllostylon rhamnoides, general view in superior focus (100x).

B: General view in optical section (100x). C: Eryngium elegans, equatorial view in optical section (100x).

D: Cissus verticillata, equatorial view in optical section (100x). E: Equatorial view in superior focus (100x).

F: Equatorial view where apertures are appreciated (100x). G: Anthoceros aﬀ. Lamellatus, polar view in

optical section (100x). H: Polar view where the trilete is appreciated (100x). I: Cyathea aﬀ. Multiﬂora, polar

view in superior focus (100x). J: Phlegmariurus aﬀ. Mandiocanus, polar view in superior focus (100x). K:

Phaeoceros bulbiculosus, polar view in optical section (100x). L: Phaeoceros tenuis, polar view in superior

focus (100x). M: Jamesonia ﬂexuosa, polar view in superior focus (100x). N: Pellaea sp., polar view in

optical section (100x). O: Selaginella aﬀ. Marginata, polar view where the trilete is appreciated (100x). P:

Sphagnum sp., polar view in optical section (100x). Scales= 10 µm.

subtriangular. Laesura with straight rays reaching Argentina (Morbelli, 1977; Morbelli et al., 2001)

to the equator. Exospore thin 1-2 μm thick. and from Brazil (Lorscheitter et al., 1998).

Baculate with less frequent spinose and verrucate

Ecological procedence: perennial herbaceous

ornamentation. Size: 17-37 µm. Micro-megaspores represented in Selaginella sellowii Hieron. grass of

belonging to this taxon were described from the natural vegetation of Corrientes (Fontana, 2018).

743

Bol. Soc. Argent. Bot. 57 (4) 2022

Sphagnaceae (Sphagnopsida, Sphagnales)

aquaticum and Ludwigia peploides, which integrate

Sphagnum sp. Fig. 5P.

marsh grasslands and hygrophilous communities

Morphology: trilete spore. Polar view suggesting locally humid conditions in the studied

subtriangular, with rounded angles. Rays of the area (Fernandez Pacella & Lara, 2019) (Fig. 6).

laesurae reach the equator. Exospore 2-4.5 μm

thick. Finely rugged. Size: 20-26 µm (Fuertes & 5800 and 5141 cal year BP is characterized by local

Rodríguez, 2009). taxa of Poaceae, Cyperaceae, Typha dominguensis,

A later stage of the Mid Holocene, between

Ecological procedence: moss well represented Sphagnum sp., representing a characteristic wetland

in marshes of the natural vegetation of the Iberá association. The presence of Typha suggests

System (Arbo & Tressens, 2002).

waterlogged or ﬂooded soils with slow-moving

water and poor drainage conditions (Cabrera,

1976; Carnevalli, 1994). These plant communities,

characteristic of lakes, swampy depressions and low

ﬂoodplains, would indicate for this later stage of

the Mid Holocene, sub-humid to humid conditions

dIScuSIon

Local and extra- local taxa and plant associations (Fernandez Pacella & Lara, 2019) (Fig. 6).

during the Holocene Between 5141 and 3506 cal year BP was

The ﬁrst stage of the Mid Holocene, between characterized by a prevalence of local herbaceous

990 and 5800 cal year BP is characterized mostly vegetation composed of Anthemis cotula, Conyza

6

by local taxa such as Imperata brasiliensis, pampeana, Baccharis trimera, Gutierrezia resinosa,

Echinochloa crus-galli, Echinochloa polystachya, Bidens subalternans, Gaillardia megapotamica,

Hymenachne pernambucensis, Cyperus rotundus, Senecio bonariensis, Amaranthus muricatus,

Crinum americanum, Cypella herbertii, Empetrum Eryngium elegans and Poaceae, characteristic of

aﬀ. rubrum, Polygala leptocaulis, Myriophyllum psammophilous herbaceous steppe (Fig. 6). The

Fig. 6. Floristic representation of local and extra-local taxa and plant associations during the Holocene.

LTmg-hc: local taxa-marsh grasslands and hygrophilous communities. LTwa: local taxa-wetland

association. LTphs: local taxa-psammophilous herbaceous steppe. LThf: local taxa-hygrophilous forest.

ExT: extra-local taxa.

744

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

presence of Chenopodiaceae suggests periodic of the understory vegetation. The interpretation of

desiccation of the water bodies (Tonello & Prieto, “humid forests community” is clearly supported by

2

008). These results suggest dry environmental the abundance of epiphytes and creeping taxa giving

conditions (Fernandez Pacella et al., 2011; Fernandez an idea of a jungle type environment (Carnevalli,

Pacella & Lara, 2019). During this stage, diﬀerent 1994; Fontana, 2018).

extra-local species are documented, which indicate

the beginning of the development of various types Comparison with modern vegetation

of forests known today as Forests of Schinopsis

All the species described herein are still part of

balansaeandMyracrodruonbalansae(“quebrachal”) the ﬂora of Corrientes. For example, Selaginella

of the humid Chaco phytogeographic region, better aﬀ. marginata present in the Mburucuyá National

represented in Chaco, Formosa and northern Santa Park (Meza Torres et al., 2013) constitutes the

Fe provinces, with impoverished remains in the NW herbaceous community of “Selaginella sellowii

of Corrientes (Fontana, 2018). The main species grass” that grows on eroded soils in the open forests

documented herein are Anadenanthera colubrina, of Schinopsis balansae and Myracrodruon balansae

Myracrodruon balansae, Calliandra parvifolia, of north-western Corrientes (Bauni & Homberg,

Myrcianthes pungens, Phyllostylon rhamnoides, 2015; Fontana, 2018). These forests characterized by

Pisonia aculeata, Prosopis alba, Schinopsis tree and shrubby forms herein documented as well

balansae, Schinus longifolia, Trichilia elegans, (Anadenanthera colubrina, Myracrodruon balansae,

Justicia brasiliana and Serjania perulacea. This type Calliandra parvifolia, Myrcianthes pungens,

of forest develops in highest topographic areas free Phyllostylon rhamnoides, Pisonia aculeata, Prosopis

of ﬂoods characterized by soils with lower moisture alba, Schinopsis balansae, Trichilia elegans, Justicia

gradients (well-drained) and a higher proportion brasiliana, Serjania perulacea).

of fine particles (Maldonado & Hohne, 2006;

Syagrus romanzoﬃana and Schinus longifolia

Morello & Rodríguez, 2009). Among these taxa, represent the extensive palm groves particularly on

Anadenanthera was recorded from the Miocene to the western coast of the Paraná River in the NW of

Pliocene in Patagonia and its last record is extended Corrientes. They develop in ﬂoodplain valleys of

into the Holocene of the Iberá Wetland (Fernandez some streams and depressed areas with soils bearing

Pacella, 2015).

a high content of ﬁne materials (silt and clays), mixed

From 3484 cal year BP until the present (Late with sand, brought by the river, generally subjected

Holocene) the palynologic record shows a prevalence to frequent, non-permanent ﬂoods (Fontana, 2018).

of herbaceous local vegetation composed of marshy

The local herbaceous taxa Imperata brasiliensis,

taxa and wetland association (Fig. 6). Local taxa Echinochloa crus-galli, Echinochloa polystachya,

from the “Hygrophilous Forest”, represented by Hymenachne pernambucense, Cyperus

Senegalia bonariensis, Celtis iguanaea, Polygonum rotundus, Eryngium elegans, Cypella herbertii,

acuminatum, Polygonum convolvulus, Cissus Myriophyllum aquaticum, Empetrum aﬀ. rubrum,

verticillata, Plinia rivularis, Tecoma stans, Polygala leptocaulis, Ludwigia peploides, Typha

Thinouia mucronata, Chrysophyllum marginatum, dominguensis, Celtis iguanaea, Anthemis cotula,

Mikania cordifolia, Cyathea atrovirens, Jamesonia Conyza pampeana, Baccharis trimera, Mikania

ﬂexuosa, Phlegmariurus mandiocanus, Pellaea sp., cordifolia, Gutierreziaresinosa, Bidenssubalternans,

Phaeoceros tenuis and Phaeoceros bulbiculosus Gaillardia megapotamica, Senecio bonariensis,

also documented in variable frequency (Fernandez Amaranthus muricatus and Sphagnum sp., are

Pacella et al., 2011; Fernandez Pacella & Lara, currently part of the hygrophilous communities of

2019). This Hygrophilous Forest developed in this marsh- grasslands, wetlands and psammophilous

interval, forms a strip that extends along small herbaceous steppe of the Iberá Wetland (Arbo &

ﬂoodplain valleys close to “madrejones” or ponds Tressens, 2002).

with reservoirs, sometimes superimposed on the

Senegalia bonariensis, Polygonum acuminatum,

grasslands (Eskuche, 2004; Garcia et al., 2013). Polygonum convolvulus, Cissus verticillata, Plinia

It is characterized by low to medium trees in less rivularis, Tecoma stans, Thinouia mucronata,

dense forests that allows enough light input for the Chrysophyllum marginatum, Cyathea atrovirens,

establishment of herbaceous communities as part Jamesonia flexuosa, Phlegmariurus saururus,

745

Bol. Soc. Argent. Bot. 57 (4) 2022

Pellaea sp., Phaeoceros tenuis and Phaeoceros acKnowledgementS

bulbiculosus, represent varied habits from tree

to epiphyte of the Hygrophilous Forests in the

The authors wish to express their gratitude to

W, N and center of Corrientes (Carnevalli, 1994; Prof. Oscar Canteros performing the chemical

Fontana, 2018).

processing of samples, to Teodoro Roberto

technical) accompanying us in the different

journeys.

Funding: This work was supported by Secretaría

General de Ciencia y Técnica, Universidad

(

concluSIon

The 55 pollen and spore types described and Nacional del Nordeste (PI 18Q006).

illustrated in this contribution reaﬃrm diversiﬁed

ﬂoras existed during the Mid-Late Holocene in

Corrientes. Of them, 46 pollen grains correspond bIblIograPhy

to 27 families of angiosperms and 9 trilete

spore-types of ferns, lycophytes and bryophytes, ACEVEDO, T. L. & L. M. ANZÓTEGUI. 1998.

obtained from six lakes of this wetland. Forty-

two (42) taxa of these lived in the Iberá wetland

Myrtaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

& G. A. CUADRADO (eds.), Flora Polínica del

Nordeste Argentino, vol. I: 67-80. EUDENE,

Corrientes.

(

local taxa), whereas 13 extra-local ones would

have arrived chieﬂy through air currents. This

ecologic/procedence differentiation of species

and their relative frequency per sample in each

core (pollen diagrams) presented by Fernandez

Pacella et al. (2011), Fernandez Pacella (2013),

Fernandez Pacella & Lara (2019) was applied

to interpret environmental characteristics and

the evolution of the Iberá vegetation. Therefore,

the identiﬁcation of pollen grains up to species

level enhances paleoenvironmental reconstructions

based on more accurate ecological information and

geographic distribution. Some local species as,

ALONSO, U. C. 2014. Palinoteca de Plantas

Vasculares Acuáticas para el Análisis Polínico en

Paleolimnología. Graduate Thesis. Universidade

da Coruña, España.

ANTON, A. M. & F. O. ZULOAGA (dir). Flora

Argentina [online]. Available in: http://www.

ﬂoraargentina.edu.ar [Access: 25 October 2021].

ANZÓTEGUI, L. M. 2001. Anacardiaceae. In: PIRE,

S. M., L. M. ANZÓTEGUI & G. A. CUADRADO

(eds.), Flora Polínica del Nordeste Argentino, vol.

II: 19-26. EUDENE, Corrientes.

considered in this studied region, were temperate- ANZÓTEGUI, L. M. & S. S. GARRALLA. 1985.

environmentally restricted and, for example, during

the ﬁrst stage of the Late Holocene, characteristic

species of psammophilous herbaceous steppe were

documented. On the other hand, in a later stage

Estudio palinológico de la Formación Paraná

(Mioceno superior), (Pozo Joseﬁna), Provincia

de Santa Fe, Argentina. I parte: Descripciones

sistemáticas. Facena 6: 101-177.

of the Late Holocene, several species appeared ANZÓTEGUI, L. M. & M. S. FERUCCI. 1998.

indicating the beginning of the development of

Hygrophilous Forest”. Instead, other species such

as grasses are dominant throughout the Holocene

in Corrientes. This work broadens the knowledge

Sapindaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

& G. A. CUADRADO (eds.), Flora Polínica del

Nordeste Argentino, vol. I: 95-110. EUDENE,

Corrientes.

“

of the palynological ﬂora of northeastern Argentine ANZÓTEGUI, L. M. & M. A. CACCAVARI. 2001.

and, will help to diﬀerentiate the local vegetation

from the extra-local in future interpretations.

Vitaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

& G. A. CUADRADO (eds.), Flora Polínica del

Nordeste Argentino, vol: 141-147. EUDENE,

Corrientes.

authorS contrIbutIon

ANZÓTEGUI, L. M. & L. R. MAUTINO. 2001a.

Celtidaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

& G. A. CUADRADO (eds.), Flora Polínica del

Nordeste Argentino, vol. II: 61-64. EUDENE,

Corrientes

Both authors have jointly and equally carried out

the data collection, interpretation and writing of the

manuscript.

746

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

ANZÓTEGUI, L. M. & L. R. MAUTINO. 2001b.

Ulmaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

G. A. CUADRADO (eds.), Flora Polínica del

Flora Polínica del Nordeste Argentino, vol. III: 55-

69. EUDENE, Corrientes.

CARNEVALLI, R. 1994. Fitogeografía de la Provincia

de Corrientes. LITOCOLOR, Asunción.

&

Nordeste Argentino, vol. II: 137-139. EUDENE,

Corrientes.

CECOTTI ÁLVAREZ, M. D., M. E. GARCÍA, N. J.

F. REYES y A. C. SLANIS. 2017. Morfología

polínica de las especies de Ludwigia (Onagraceae,

Ludwigioideae) del Noroeste de Argentina. Lilloa

54: 29-40.

CONTRERAS-DUARTE, A. R., R. G. BOGOTÁ-

ÁNGEL & L. C. JIMÉNEZ-BULLA. 2006.

Atlas de las Esporas de Pteridóﬁtos de Chipaque

(Cundinamarca, Colombia). Caldasia 28: 327-357.

CUADRADO, G. A. 1998a. Polygalaceae. In: PIRE, S.

M., L. M. ANZÓTEGUI & G. A. CUADRADO

(eds.), Flora Polínica del Nordeste Argentino, vol. I:

81-88. EUDENE. Corrientes.

APG IV. 2017. Angiosperm Phylogeny Group

classiﬁcation for the orders and families of ﬂowering

plants. Checklist dataset [online]. Available in:

https://doi.org/10.15468/fzuaam accessed via GBIF.

org

ARBO, M. & S. TRESSENS. 2002. Flora del Iberá:

Corrientes, Argentina. EUDENE, Corrientes.

ARREGUÍN-SÁNCHEZ, M., R. FERNÁNDEZ NAVA,

R. PALACIOS CHÁVEZ & D. L. QUIROZ

GARCÍA. 1996. Morfología de las esporas de

Pteridóﬁtas Isospóreas del Estado de Querétaro,

México (parte a). Polibotánica 2: 10-60.

BAUERMANN, S. G., A. CARDOSO PACHECO

EVALDT, J. R. ZANCHIN & S. A. DE LORETO

BORDIGNON. 2010. Diferenciação polínica de

Butia, Euterpe, Geonoma, Syagrus e Thritrinax e

implicações paleoecológicas de Arecaceae para o

Rio Grande do Sul. IHERINGIA, Sér. Bot. 65: 35-

CUADRADO, G. A. 1998b. Sapotaceae. In: PIRE, S. M.,

L. M. ANZÓTEGUI & G. A. CUADRADO (eds.),

Flora Polínica del Nordeste Argentino, vol. I: 111-

117. EUDENE. Corrientes.

CUADRADO, G. A. 1998c. Polygalaceae. In: PIRE, S.

M., L. M. ANZÓTEGUI & G. A. CUADRADO

(eds.), Flora Polínica del Nordeste Argentino, vol. I:

81-88. EUDENE, Corrientes.

4

6. https://isb.emnuvens.com.br/iheringia/article/

view/79

BAUNI, V. & M. A. HOMBERG. 2015. Reserva Natural

Campo San Juan. 1a ed. Fundación de Historia

Natural Félix de Azara, Buenos Aires.

BHATTACHARYA, K., M. R. MAJUMDAR & S.

GUPTA BHATTACHARYA. 2009. A Text Book of

Palynology. New Central Book Agency (P) Ltd.,

Kolkata.

BOELCKE, O. 1992. Plantas Vasculares de la Argentina:

nativas y exóticas. 2da ed. Hemisferio Sur, Buenos

Aires.

CABRAL, E. L. & M. CASTRO. 2007. Palmeras

Argentinas, Guía para el reconocimiento. L.O.L.A.,

Buenos Aires.

CABRERA, A. L. 1976. Regiones fitogeográficas

argentinas. En: Kugler, W. F. (ed.), Enciclopedia

argentina de agricultura y jardinería. Tomo 2. 2a.

edición, Fascículo 1. pp. 1-85. Acme, Buenos Aires.

CACCAVARI, M. A. & E. DOME. 2001. Fabaceae-

Mimosoideae. Tribu: Acacieae. In: PIRE, S. M.,

L. M. ANZÓTEGUI & G. A. CUADRADO (eds.),

Flora Polínica del Nordeste Argentino, vol. II: 65-

CUADRADO, G. A. & J. J. NEIFF. 1993. Palynology

of embalsados in distrophic lakes in northeastern of

Argentina. Revista Brasil. Biol. 53: 443-451.

CURTIS, H., N. S. BARNES, A. SCHNEK & G.

FLORES. 2001. Biología. 6a. ed. en español.

Panamericana, Buenos Aires.

DELLA,A.P.&J.PRADO.2020.Jamesonia(Pteridaceae)

in Brazil. Biota. Neotrop. 20: e20200986. https://doi.

org/10.1590/1676-0611-BN-2020-0986

DELLA, A. P., J. PRADO & R. Y. HIRAI. Jamesonia. In:

Flora do Brasil [online]. Available in: http://reﬂora.

jbrj.gov.br/reﬂora/ﬂoradobrasil/FB134680

DÍEZ DAPENA, M. J., S. TALAVERA LOZANO y P.

GARCÍA MURILLO. 1988. Contributions to the

palynology of hydrophytic, non-entomophilous

angiosperms. 1. Studies with LM and SEM.

Candollea 43: 147-158.

ESKUCHE, U. G. 2004. La vegetación de la vega del

rio Paraná medio superior, Argentina. Folia Bot.

Geobot. Correntesiana 17:1-60.

FAEGRI, K. & D. IVERSEN. 1989. Texbook of pollen

analysis. 4 ed. New John Willey & Sons, Chichester.

FERNÁNDEZ, I. 1987. Contribución al conocimiento

palinológico de Cyperaceae. Acta Bot. Malac. 12:

173-181.

7

2. EUDENE, Corrientes.

CACCAVARI, M. A. & E. DOME. 2006. Fabaceae-

Mimosoideae. Tribu: Mimoseae. In: PIRE, S. M.,

L. M. ANZÓTEGUI & G. A. CUADRADO (eds.),

747

Bol. Soc. Argent. Bot. 57 (4) 2022

FERNANDEZ PACELLA, L. 2013. Palinología del

Cuaternario en sedimentos lacustres del Noroeste

del Iberá, Corrientes, Argentina. Ph.D. Thesis.

Universidad Nacional del Nordeste, Argentina.

FERNANDEZ PACELLA, L. 2014. Morfología polínica

de especies del género Senna (Fabaceae) del sureste

del Iberá, Corrientes, Argentina. Rev. Biol. Trop. 62:

FUERTES, E. & M. RODRÍGUEZ. 2009. Estudio sobre

la morfología y germinación de las esporas de tres

species de Sphagnum (Murci, Sphagnaceae). Bot.

Complut. 33: 29-35. https://revistas.ucm.es/index.

php/BOCM/article/view/BOCM0909110029A

GARCÍA, A. V., H. S. LEYES, R. B. MARTÍNEZ, Y.

J. PÉREZ, J. M. PINEIRO, M. E. PRIETO & S.

C. SCHALLER. 2013. Guía de la Vegetación de la

Estación Biológica Corrientes. In: FONTANA, J.

L. (ed.), Publicaciones Didácticas de la Cátedra de

Ecología Vegetal, vol. I: 44-51. Facultad de Ciencias

Exactas, Naturales y Agrimensura, Corrientes.

GARRALLA, S. S. 1998. Estudio palinológico de una

secuencia sedimentaria del Holoceno, norte de Santa

Fe, Argentina. Polen 9: 17-27.

GARRALLA, S. S. & G. A. CUADRADO. 2001.

Meliaceae. In: PIRE, S. M., L. M. ANZÓTEGUI

& G. A. CUADRADO (eds.), Flora Polínica del

Nordeste Argentino, vol. II: 95-100. EUDENE,

Corrientes.

GENTILI, C. & RIMOLDI, H. 1979. Mesopotamia. II

Simposio de Geología RegionalArgentina. Academia

Nacional de Ciencias en Córdoba, Vol. I: pp 185-222.

GÓMEZ-NOGUEZ, F., B. PÉREZ-GARCÍA, A.

MENDOZA-RUÍZ & A. OROZCO-SEGOVIA.

2013. Flora Palinológica de los Helechos y Licoﬁtas

de Río Malila, Hidalgo, México. Bot. sci. 91: 135-

154. https://doi.org/10.17129/botsci.410

7

69-782.

FERNANDEZ PACELLA, L. 2015. Registro fósil y

distribución de Anadenathera en Argentina desde el

Mioceno hasta la actualidad. Estud. Geol. 71: e031.

http://dx.doi.org/10.3989/egeol.41834.343

FERNANDEZ PACELLA, L. 2018. Estudio palinológico

de un perﬁl sedimentario del Holoceno Medio-

Tardío, oeste del Iberá, Corrientes, Argentina. Rev.

Mex. Cienc. Geol. 35: 93-101.

https://doi.org/10.22201/cgeo.20072902e.2018.1.531

FERNANDEZ PACELLA, L. & O. CANTEROS.

2

014. Poaceae. Tribus: Andropogoneae, Bromeae,

Cynodonteae, Eragrostidae, Olyreae y Oryzeae.

In: PIRE, S. M., L. M. ANZÓTEGUI & G. A.

CUADRADO (eds.), Flora Polínica del Nordeste

Argentino, vol. IV: 88-94. EUDENE, Corrientes.

FERNANDEZ PACELLA, L. & M. B. LARA. 2019.

Paleoenvironmental Interpretation of the mid‐late

Holocene of Corrientes province, Argentina. Nordic J.

Bot. 2019: e02252. http://dx.doi.org/10.1111/njb.02252

FERNANDEZ PACELLA, L., S. S. GARRALLA & L.

M. ANZÓTEGUI. 2011. Cambios de la vegetación

durante el Holoceno en la región Norte del Iberá.

Provincia de Corrientes, Argentina. Rev. Biol. Trop.

GRADSTEIN, S. R. 2018. Key to hornworts

(Anthocerotophyta) of Colombia. Caldasia 40:

262-270.

5

9: 103-112.

FERNANDEZ PACELLA, L., L. M. ANZÓTEGUI

Y. HORN. 2014a. Fabaceae-Caesalpinioideae.

https://dx.doi.org/10.1544 6/caldasiav40n2.71750

HERBST, R. & SANTA CRUZ, J. 1985. Mapa

litoestratigráfico de la Provincia de Corrientes.

Revista D’Orbignyana 2: 1-51.

HIRAI, R. Y. & J. PRADO. Pellaea. In: Flora do Brasil

[online]. Available in: http://reflora.jbrj.gov.br/

reﬂora/ﬂoradobrasil/FB91954 [Access: 2 November

2021].

IRIONDO, M. 1994. Los climas cuaternarios de la región

Pampeana. Museo Provincial de Ciencias Naturales

“Florentino Ameghino” 4: 481.

IRIONDO, M. 2010. Geología del Cuaternario en

Argentina. Editorial Moglia, Corrientes.

KREMP, G. W. 1965. Morphologic Encyclopedia of

Palynology. The University Arizona Press, Tucson.

LORSCHEITTER, M. L.,A. R.ASHRAF, R. MACHADO

BUENO & V. MOSBRUGGER. 1998. Pteridophyte

spores of Rio Grande do Sul ﬂora, Brazil. Part I.

Palaeontographica 246: 1-113.

&

In: PIRE, S. M., L. M. ANZÓTEGUI & G. A.

CUADRADO (eds.), Flora Polínica del Nordeste

Argentino, vol. IV: 61-77. EUDENE, Corrientes.

FERNANDEZ PACELLA, L., L. M. ANZÓTEGUI &

L. R. MAUTINO. 2014b. Fabaceae-Mimosoideae.

Tribu: Mimoseae-Prosopis In: PIRE, S. M., L. M.

ANZÓTEGUI & G. A. CUADRADO (eds.), Flora

Polínica del Nordeste Argentino, vol. IV: 88-94.

EUDENE, Corrientes.

FLORA DO BRASIL [online]. Available in: http://

ﬂoradobrasil.jbrj.gov.br/ [Access: 25 October 2021].

FONTANA, J. L. 2018. La Vegetación del Nordeste

Argentino. 1. Las Comunidades vegetales del

Noroeste de Corrientes y del Este de Chaco.

Publicaciones Didácticas de la Cátedra de Ecología

Vegetal. Vol. 3. UNNE, Corrientes.

748

L. Fernandez Pacella & M. Di Pasquo - Pollen and spores of the Iberá Wetlands

MALDONADO, P. & E. HOHNE. 2006. Atlas del Gran

PEÑALOZA-BOJACÁ, G. F., B. A., OLIVEIRA,

C. A. T. ARAUJO, L. B. FANTECELLE, J. C.

VILLARREAL & A. S. MACIEL-SILVA.

Anthocerotaceae. In: Flora do Brasil [online].

Available in: http://reflora.jbrj.gov.br/reflora/

ﬂoradobrasil/FB97153 [Access: 3 April 2021]

PÉREZ-JIMÉNEZ, J. C., F. ESLAVA-SILVA, K.

JIMÉNEZ-DURÁN, F. GÓMEZ-NOGUEZ &

M. E. MUÑIZ-DIAZ DE LEÓN. 2020. Estudio

Palinológico de los helechos y licoﬁtas de la Zona

Núcleo Poniente de la Reserva Ecológica del

Pedregal de San Ángel, Ciudad de México, México.

Bot. Sci. 98: 517-532.

a.

Chaco americano. 1 ed. Agencia Alemana de

Cooperación Técnica, Buenos Aires.

MARKGRAF, V. & H. L. D`ANTONI. 1978. Polen

Flora of Argentina-Modern Spore and Pollen Types

of Pteridophyta, Gymnospermae and Angiospermae.

The University of Arizona Press, Tucson.

MEDEANIC, S., C. V. CORDAZZO & L. G. LIMA.

2

008. Diversidade Polínica de Plantas em Dunas do

Extremo Sul do Brasil. GRAVEL 6: 67-80. Available

in: http://repositorio.furg.br/handle/1/1867

MEZA TORRES, E. I., E. R. DE LA SOTA & M.

S. FERRUCCI. 2013. Sinopsis de los helechos

y licofitos del Parque Nacional Mburucuyá

https://doi.org/10.17129/botsci.2424

(

Corrientes, Argentina). Claves de especies. Bol.

PIRE, S. M., L. M. ANZÓTEGUI, L. R. MAUTINO &

S. S. GARRALLA. 2006. Acanthaceae. In: PIRE,

S. M., L. M. ANZÓTEGUI & G. A. CUADRADO

(eds.), Flora Polínica del Nordeste Argentino, vol.

III: 15-38. EUDENE, Corrientes.

PPG I. 2016. The Pteridophyte Phylogeny Group.

A community-derived classification for extant

lycophytes and ferns. J. Syst. Evol. 54: 563-603.

PRIETO, A. R. & M. E. QUATTROCCHIO. 1993.

Brióﬁtas y Pteridóﬁtas en sedimentos del Holoceno

en la provincia de Buenos Aires, Argentina. An.

Asoc. Palinol. Leng. Esp. 6: 17-37.

Soc. Argent. Bot. 48: 121-136.

MORBELLI, M. A. 1977. Esporas de las especies

argentinas de Selaginella (Selaginellaceae–

Pteridophyta). Obra Cent. Mus. La Plata 3: 121-150.

MORBELLI, M. A., J. R. ROWLEY & D. CLAUGHER.

2

001. Spore wall structure in Selaginella (Lycophyta)

species growing in Argentina. Bol. Soc. Argent. Bot.

6: 315-368.

MORBELLI, M. A., M. R. PIÑEIRO & G. E. GIUDICE.

010. Spore morphology and wall ultrastructure of

3

2

Hymenophyllaceae Link (Pteridophyta) from north-

west Argentina. Grana 49: 37-46.

PUNT, W., P. P. HOEN, S. BLACKMORE, S. NILSSON

& A. LE THOMAS. 2007. Glossary of pollen and

spore terminology. Rev. Palaeobot. Palynol. 143:

1-81.

http://dx.doi.org/10.1080/00173130903483663

MORELLO, J. H. &A. F. RODRÍGUEZ. 2009. El Chaco Sin

a.

Bosques. 1 ed. Ed. Orientación Gráﬁca, Buenos Aires.

NEIFF, J. J. 1997. Ecología evolutiva del macrosistema

Iberá (Corrientes, Argentina). Master Thesis.

Universidad Nacional del Litoral, Argentina.

NEIFF, J. J. 2004. El Iberá… ¿en peligro? Fundación

Vida Silvestre, Buenos Aires.

NILSON, S. & J. PRAGLOWSKI. 1992. Erdtman`s

handook of palynology. Munksgaard International

Publishers, Copenhagen.

OLIVEIRA, P. P. & F. RIBEIRO DO SANTOS. 2014.

Prospecçao palinológica em méis da Bahia. Mídia

Editora, Bahia.

ORFEO, O. 2005. Historia Geológica del Iberá, provincia

de Corrientes, como escenario de biodiversidad.

INSUGEO Miscelánea 14: 71-78.

ORFEO, O. & J. J. NEIFF. 2008. Esteros del Iberá: un

enorme laboratorio a cielo abierto. Sitios de interés

geológico de la República Argentina 46: 415-425.

PALDAT, A palynological database [online]. Available

in: (2000 onwards, www.paldat.org) [Access: 5

November 2021].

https://doi.org/10.1016/j.revpalbo.2006.06.008

RAMOS, V. 1999. Las provincias geológicas del

territorio argentino. Instituto de Geología y Recursos

Minerales. 29: 41-96.

RAVEN, P. H., R. F. EVERT & S. E. EICHHON. 1991.

Biología de las Plantas. T. I-II. Reverté, Barcelona.

RINCÓN BARON, E. J., C. H. ROLLERI, L. M.

PASSARELLI, S. ESPINOSA MATÍAS & A.

M. TORRES. 2014. Esporogénesis, esporodermo

y

ornamentación de esporas maduras en

Lycopodiaceae. Rev. Biol. Trop. 62: 1161-1195.

SÁENZ LAÍN, C. 2004. Glosario de términos

palinológicos. Lazaroa 25: 93-112.

SALGADO, C. R., 2006. Flora melífera de la provincia

de Chaco. PROSAP y Ministerio de Producción del

Chaco, Resistencia.

SERSIC, A. N. 1983. Ontogenia del esporangio

y esporogénesis en Lycopodium saururus

(Lycopodiales). Bol. Soc. Argent. Bot. 22: 205-

220.

749

Bol. Soc. Argent. Bot. 57 (4) 2022

TONELLO, M. S. & A. R. PRIETO. 2010: Tendencias

climáticas para los pastizales pampeanos durante

el Pleistoceno tardío-Holoceno: estimaciones

cuantitativas basadas en secuencias polínicas fósiles.

Ameghiniana 7: 501-514. https://www.ameghiniana.

org.ar/index.php/ameghiniana/article/view/279

TRIGO, M. & I. FERNÁNDEZ. 1995. Contribución

al estudio polínico de especies ornamentales

con interés alergógeno en cultivadas en Málaga:

Monocotiledóneas. Acta Bot. Malac. 20: 61-70.

Available in: http://ﬂoradobrasil.jbrj.gov.br/jabot/

ﬂoradobrasil/FB128519 [Access: 19 August 2021]

YBERT, J. P., R. SCHEEL-YBERT & M. CARVALHO.

2016. Grãos de pólen de plantas vasculares

Dicotiledôneas do Estado do Rio de Janeiro, Brasil.

Vol. I. Museu Nacional, Universidade Federal do

Rio de Janeiro, Rio de Janeiro.

YBERT, J. P., M. CARVALHO & R. SCHEEL-YBERT.

2018. Grãos de pólen de plantas vasculares do

Estado do Rio de Janeiro, Brasil. Vol. IV. Museu

Nacional, Universidade Federal do Rio de Janeiro,

Rio de Janeiro.

WINDISCH, P.G., C. G. V. RAMOS & B. OELLGAARD.

Lycopodiaceae. In: Flora do Brasil [online].

750