Studying Pollen rePreSentation of vegetation

and Plant richneSS from PamPean coaStal duneS

argentina, South america): exPloring from local to

landScaPe quantitative linKageS

(

eStudio de la rePreSentación del Polen de la vegetación y la riqueza de

PlantaS de laS dunaS coSteraS PamPeanaS (argentina, américa del Sur):

exPlorando laS relacioneS cuantitativaS deSde lo local al PaiSaje

1

1,2

1

Carolina Vásquez , Gonzalo Sottile * , Víctor Merino-Campos

&

1

Silvina Stutz

Summary

Background and aims: Studying plant dynamics through past pollen records may

contribute to a better understanding of long-term changes in plant communities.

Thus, this study aims to establish whether surface pollen composition and richness

of the Argentinean Coastal Dune System reﬂect landscape heterogeneity in coastal

dune environments.

1

. Grupo de Paleoecología y

Palinología, Facultad de Ciencias

Exactas y Naturales, Instituto de

Investigaciones Marinas y Costeras

(Universidad Nacional de Mar del

M&M: Twenty-four sediment surface samples were collected in small lagoons.

Landscape-scale heterogeneity up to 2000 m was mapped and classiﬁed in

landscape units. Multivariate analyses were used to classify pollen samples and

compare them to landscape unit coverage (%). Also, we evaluate plant species and

pollen richness relationship by linear regression models.

Results: The relationship between plant and pollen richness is inﬂuenced by

taxonomic smoothing, pollen production and taphonomic constraints (dispersal and

preservation). The pollen assemblages and pollen richness from surface sediments

of small lagoons and interdune slacks reﬂect plant richness and vegetation

heterogeneity at the landscape scale (ca. 1000-2000 m). The main contributors

to pollen richness are anemophilous pollen types, although some entomophilous

pollen types are useful to infer some local heterogeneity.

Plata - CONICET), Mar del Plata,

Argentina

2

. Grupo de Biología y Ecodiversidad

vegetal, Facultad de Ciencias

Exactas y Naturales, Instituto de

Investigaciones Marinas y Costeras

(

Universidad Nacional de Mar del

Plata - CONICET), Mar del Plata,

Argentina

*gonzalo_sottile@yahoo.com.ar

Conclusions: We report the ﬁrst quantitative analysis on pollen-vegetation

relationship of coastal ecosystems showing that pollen records reﬂect landscape

vegetation attributes, encouraging the study of past plant diversity and landscape

variability based on pollen records.

Citar este artículo

VÁSQUEZ, C., G. SOTTILE, V.

MERINO-CAMPOS& S. STUTZ. 2024.

Studying pollen representation of

vegetation and plant richness from

Pampean coastal dunes (Argentina,

South America): exploring from

local to landscape quantitative

linkages. Bol. Soc. Argent. Bot. 59:

Key wordS

Coastal dune vegetation, landscape heterogeneity, pollen, quantitative pollen-

vegetation relationship.

reSumen

5

07-527.

Antecedentes y objetivos: Estudiar la dinámica de la diversidad vegetal mediante

registros polínicos puede contribuir a entender patrones de cambio de largo plazo.

Para ello es necesario estudiar cómo la composición y riqueza del polen superﬁcial

de los sistemas de Dunas Costeros argentinos reﬂejan la heterogeneidad del paisaje.

M&M: Se recolectaron 24 muestras de superﬁcie de sedimentos de pequeñas

lagunas. Se mapeó y clasiﬁcó la heterogeneidad vegetal hasta 2000 m en

unidades de paisaje y mediante análisis multivariados y agrupamiento se comparó

el contenido polínico y la cobertura de las unidades de paisaje (%). Se modeló la

relación entre riqueza especíﬁca y riqueza polínica, desde la escala local hasta la

de paisaje mediante regresiones lineales.

DOI: https://doi.

org/10.31055/1851.2372.v59.

n4.44793

Resultados: La relación entre la riqueza especíﬁca y polínica está inﬂuenciada por

el sesgo taxonómico, la producción polínica y limitaciones tafonómicas (dispersión

y preservación de polen). Las asociaciones y riqueza de polen superﬁcial reﬂejan la

riqueza especíﬁca y heterogeneidad vegetal a escala de paisaje (ca. 1000-2000 m).

Los principales contribuyentes a la riqueza polínica son anemóﬁlos. Algunos tipos

de polen entomóﬁlos son útiles para inferir cierta heterogeneidad local.

Recibido: 17 Abr 2024

Aceptado: 24 Jul 2024

Publicado en línea: 31 Dic 2024

Publicado impreso: 31 Dic 2024

Editor: Gonzalo J. Márquez

Conclusiones: El modelado cuantitativo de la relación polen-vegetación en término

de unidades de paisaje y riqueza polínica demuestra el alto potencial para la

aplicación de estos modelos en la reconstrucción de la dinámica de la vegetación

en este tipo de ambientes a partir de registros polínicos.

PalabraS clave

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

Heterogeneidad del paisaje, polen, relación cuantitativa polen-vegetación, vegetación

de dunas costeras.

507

Bol. Soc. Argent. Bot. 59 (4) 2024

introduction

surface samples (e.g. Goring et al., 2013; Meltsov

et al., 2011; Matthias et al., 2015; Felde et al., 2015;

Coastal Dune Systems (CDS) are landforms of Gosling et al., 2018; Reitalu et al., 2019; Abraham

regional extension, consisting mainly of mobile et al., 2020; Papadopoulou et al., 2022).

dunes and dune ridges ﬁxed by vegetation, in which

Modern pollen-vegetation studies in Argentinean

a mosaic of environments has been formed with CDShavebeencarriedoutonthecoastofSanMatías

plant communities adapted to diﬀerent landforms. Gulf (north Patagonia) (Marcos & Mancini, 2012),

They are abundant on the Argentinian coast, and and in the southwest and southeast coast of Buenos

according to Isla et al. (1996, 2001) and Codignotto Aires Province (Fontana, 2003, 2004, 2005b; Stutz

et al. (2012) developed during the late Holocene & Prieto, 2003; Latorre et al., 2010; Monserrat et

between ca. 3070 and 540 yrs. BP. These systems al., 2012). These studies mainly focused on the

are highly dynamic and diverse, in terms of relationship between vegetation distribution and

vegetation cover and composition. They include pollen dispersal. Fontana (2005a) emphasized

endemic species and provide multiple ecosystem that pollen representation was inﬂuenced both by

services of economic and environmental value diﬀerences in pollen production, dispersal, and

Celsi (2016). Since the XIX century, they have preservation of individual taxa, as well as by the

been threatened by several factors, including spatial distribution of vegetation and topography of

aﬀorestation with exotic species of Acacia Mill., the CDS and by wind pattern.

Populus L., Eucalyptus L'Hér., and Pinus L. genera,

Since the pioneers of quantitative analysis of

which dispersion now represents one of the main modern and past pollen-vegetation relationships,

threats to conservation in the area. Invasions of Prentice (1985), Sugita (1994, 2007a, b) and Bunting

woody species have been considered to affect & Middleton (2005, 2009), many works have

severely large areas of grassland causing variations been carried out, mainly in northern hemisphere

in the dominant life forms, disturbing the dynamics forests and steppe ecosystems. Pollen richness or

of vegetation and changing the distribution patterns rareﬁed palynological richness was ﬁrst interpreted

of species and increasing ecosystem patchiness as a proxy of past plant diversity (e.g. Birks &

(Alberio & Comparatore, 2014). Invasion rates and Line, 1992; Bennett et al., 1992; Bunting, 1994;

its impacts on biodiversity have been characterized Andersen, 1995; Giesecke et al., 2012), however,

spatially and seasonally, nevertheless, long term during the last twenty years, some quantitative

eﬀects (decadal or longer time lags) have not been approaches have explored the relationship between

recorded for Argentinian CDS ecosystems.

modern vegetation and modern pollen assemblages

Pollen assemblages preserved in sediments of from surface samples to discuss the performance of

CDS lagoons and interdune slacks can be used these modeling and appliance to past palynological

as a proxy for past plant richness as they record records (e.g. Burry et al., 2001; Goring et al., 2013;

the dynamics of plant taxa through both, space Masciadri et al., 2013; Felde et al., 2015; Matthias et

and time. However, pollen assemblages studies al., 2015; Reitalu et al., 2019; Li et al., 2022; Senn

are limited by low taxonomic resolution, pollen et al., 2022). These studies warned about the spatial

production, and taphonomic processes (transport, scale-dependence and the ﬂoristic composition of

deposition, and preservation). Moreover, pollen- speciﬁc ecosystem eﬀects on the capacity of this

based species richness estimates are sensitive to proxy to reconstruct past plant diversity dynamics.

source area size, missing taxa, and pollen count Disentangling between natural and human forcing

sizes (Van der Sande et al. 2021) which may of past Coastal Dune System vegetation dynamics

decrease the accuracy with which pollen represents demands calibrating modern pollen-vegetation

vegetation communities (Goring et al. 2013). For representation models since a quantitative approach.

these reasons, pollen abundance cannot be directly

The CDS presents a large number of interdune

translated into plant abundance when interpreting slacks, which have a high potential for pollen

past pollen assemblages (Fontana, 2005a). To trapping and preservation, as well as a mixture of

understand fossil pollen assemblages and make landscape units with vegetated and pristine areas and

ecological inferences, it is necessary to calibrate some areas invaded by exotic forests, particularly in

modern pollen-vegetation diversity relationships in the northern section of the CDS, which makes it

508

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

an ideal area for performing paleoenvironmental while hydrophytic plants grow in ﬂooded lowlands

reconstructions of past vegetation changes (lagoons and interdune slacks) (Celsi, 2016).

(Vásquez et al., 2023). Previous studies based on

The vegetation of the CDS is characterized

fossil pollen records of Faro Querandí (northern mainly by psammophytic species that are distributed

area of the CDS) registered changes in wetland and according to the geomorphology heterogeneity

dune vegetation over the last 500 years and they (Cabrera, 1941; Stutz & Prieto, 2003; Celsi, 2016;

recorded also the ﬁrst expansion of exotic forests Marcomini et al., 2017). According to Fontana

in the CDS (Vásquez et al., 2023). However, these (2005a) this heterogeneity can be zoned in diﬀerent

pollen-based qualitative reconstructions have not landscape units. These landscape units include a strip

discussed long-term changes in plant diversity. of beach free of vegetation and the back shore, the

Thus, modeling between modern plant and pollen adjacent area with permanent sand, salt spray, and

richness and pollen-vegetation representation in a water droplets carried by the wind in storms. Only a

quantitative dimension would encourage future past few halophytic plant species (<5% coverage), such

vegetation and diversity changes in quantitative as Sporobolus coarctatus (Trin.) P.M. Peterson &

reconstructions.

Saarela, grow under such environmental conditions.

The aim of this work is to establish whether pollen Mobile dunes, active dunes with a strong substrate

composition and richness reﬂect the landscape movement due to wind action, expand toward

heterogeneity, considering both vegetation patterns the mainland, with <10-30% coverage. This

and geomorphological characteristics of the CDS. zone is characterized by pioneer species such as

We plan to evaluate the relationship between Panicum racemosum (P. Beauv.) Spreng., Calycera

plant and pollen richness in relation to pollen crassifolia (Miers) Hicken, Senecio crassiﬂorus

production, taxonomic and taphonomic (dispersal (Poir.) DC. and the exotic Cakile maritime Scop.

and preservation) constraints, at local and extra- Inward to the continent, the vegetation cover

local scales.

increases, stabilizing dunes sediments favoring

the formation of humus. These semiﬁxed (>30%

coverage) and fixed dunes (>75% coverage)

are covered by grasslands and psammophytic

shrublands dominated by Poa lanuginose Poir.,

Tessaria absinthioides (Hook. & Arn.) DC., and

materialS and methodS

Study area

The study area corresponds to the Faro Querandí Achyrocline satureioides (Lam.) DC. Herbaceous

Municipal Nature Reserve (FQNR), located in the plants such as Ambrosia tenuifolia Spreng., Adesmia

Coastal Dune Systems (CDS) of the southeast of incana Vogel, and Hydrocotyle bonariensis Lam.

Buenos Aires Province, Argentina, which preserves are subordinated components joint with some shrub

a valuable area of the Pampa plain ecoregion, species, Baccharis genistifolia DC., Margyricarpus

characterized mainly by the presence of grassland pinnatus (Lam.) Kuntze, and Discaria Americana

vegetation (Fig. 1). The CDS constitutes an almost Gillies & Hook., and the exotics Centaurium

uninterrupted extensive coastal dune ﬁelds with a pulchellum (Sw.) Druce, Blackstonia perfoliata

width, from the sea towards the mainland, ranging (L.) Huds., Medicago lupulina L., Melilotus indicus

from hundreds of meters to 8 km (Celsi, 2016). (L.) All., and M. albus Desr. The interdune slacks

These dunes are generally parabolic to pyramidal, (ca. 100% coverage) are intermediate depressions

although there are also transverse dunes, barkhans frequently ﬂooded covered by species of the genus

(

crescent-shaped), and star-shaped dunes (Bértola et Typha, other hydrophytic such as Schoenoplectus

al., 2002). Speciﬁcally, the FQNR comprises about spp. and Eleocharis spp., Juncus sp. and other

000 ha with an aeolian morphology, and dunes herbaceous dicots, Bacopa monnieri (L.) Wettst.,

5

are mainly transversal, with a general southwest- and Eryngium spp. Cortaderia selloana (Schult.

northeast orientation (Codignotto et al., 2012). The & Schult. f.) Asch. & Graebn. grow on the edges

pronounced topographic and edaphic variations at of the interdune slacks and can also form large tall

the CDS drive noticeable changes in vegetation grasslands in the semiﬁxed dunes or even patches

cover and ﬂoristic composition. Psammophytic on mobile dunes slopes. Temporary or permanent

plant types cover mobile and semiﬁxed dunes lagoons develop due to the contribution of rainwater

509

Bol. Soc. Argent. Bot. 59 (4) 2024

Fig. 1. Map of study area in Pampa plain (according to Oyarzabal et al., 2018) showing the location of

Reserve Faro Querandí and other sites mentioned in the text.

or water table raising, where dense hydrophytic

The climate is humid temperate with a mean

vegetation grows, as species of the Cyperaceae and annual temperature of 14 ºC, with marked

Juncaceae families, Potamogeton spp, Polygonum seasonality; the average yearly precipitation

punctatum Elliott, Utricularia gibba L., among is 930 mm and the mean annual humidity is

others.

greater than 75% (Estación Meteorológica

Since the early 20th century, exotic trees were Aeródromo de Villa Gesell 37º14’ S, 57º01’ W,

introduced to ﬁx mobile dunes in the area of Villa Servicio Meteorológico Nacional). The most

Gesell village and to place the Querandí lighthouse, frequent winds are those coming from the north,

like pines, eucalyptus, cypresses, poplars, acacias, but the most intense are those coming from the

tamarisks, and various fruit trees (Benseny, 2011; southeast, south, and southwest (Bértola et al.,

Provendola, 2013). As a result, at present wide 1999).

areas have been invaded mainly by Populus alba

L., Pinus spp., and Acacia spp., particularly in the Sampling and vegetation surveys

northern and central sector of the FQNR. Other

Between December 2019 and March 2023, 24

exotic trees (Betula pendula Roth, Casuarina sp., sediment surface samples, henceforth sampling

Juglans nigra L., Araucaria araucana (Molina) K. sites, were collected in lagoons and interdune

Koch and A. angustifolia (Bertol.) Kuntze, Cedrus slacks of the CDS, between 37º32’ S 57º11’ W

spp.) are widely spread across the gardens of some and 37º22’ S 57º3’ W (Fig. 2), with a spatula or

countryside residences neighboring the FQNR a Gravity-corer type sampler, depending on the

(Stutz, 2001).

presence of water during the sampling.

510

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

Fig. 2. Location of the sampling sites and surrounding landscape units of the Faro Querandí Municipal

Nature Reserve, in a 2000 m radius: A close-up of the sampling sites in the southern, central and northern

zones of the FQNR, a detail of the Landscape units of a sampling site and of scheme of the vegetation

surveys method (adapted from Bunting et al., 2013), are shown. Squares (=quadrat) shows vegetation

2

survey sampling plot (5 m ) and the distances to the central point of the circumference (central point, 10 m

and 20 m). Notice the 10 m and the 20 m radii plots are distributed following the main four cardinal directions.

To characterize the vegetation at a local scale, 5x5 m plots inside the circumference. A total of 9

concentric censuses were carried out concerning plots per sampling site, one of them placed at the

each sampling site, at 10- and 20-meter radii, center of the circumference, 4 plots placed at 10m

distributed following the four main cardinal radius and other 4 plots placed at 20 m radius were

directions (adapted from Bunting et al., 2013). measured (Fig. 2). A total of 216 vegetation surveys

According to Prentice (1985), the term ‘local’ were obtained.

refers to pollen input within 20 m of the basin edge,

To evaluate pollen representation of extra-

‘extra-local’ refers from 20 m to 2 km, ‘regional’ local vegetation, we mapped landscape scale

refers from 2 to 200 km, and ‘extra-regional’ from heterogeneity up to 2000 m radii from the sampling

beyond 200 km. Cover, as percentages, of every site. We summarized landscape scale heterogeneity

vascular plant species was measured at diﬀerent data by deﬁning ten diﬀerent landscape units,

511

Bol. Soc. Argent. Bot. 59 (4) 2024

following Fontana (2005a), vegetation and the study area or its surroundings are recorded, list

geomorphological features (see Study area elaborated according to vegetation surveys between

section), using 2020 Google Earth images by visual 2019 and 2023 and literature review (Cabrera,

interpretation, and ﬁeld observations. The identiﬁed 1941; Alberio, 2010; Benseny, 2011; Stutz, 2000;

landscape units were: exotic forest, interdune Stutz, 2001; Alberio & Comparatore, 2014; Celsi,

slacks, ﬁxed, semiﬁxed and mobile dunes, beach, 2016), about the diﬀerent pollen types recorded in

back shore, lagoons, buildings and roads, and the pollen counts. It is shown to which group each

Atlantic Ocean (Fig. 2). The area of each landscape pollen type was assigned, and data such as the habit

unit was calculated within the 1000 and 2000 and pollination type of the plant are added.

m radii (extra-local scales) using QGis software

version 3.22.11).

Pollen diagrams were plotted using TILIA

2.6.1 (Grimm, 2020). Each pollen type grouped in

Dunes and each pollen type grouped in Grasslands

was calculated as a percentage of the Dunes plus

(

Pollen analysis

Surface sediment samples were processed Grasslands pollen sum (∑D+G); each pollen type

following the standard protocol for palynomorph grouped in Wetlands were calculated as a percentage

extraction(Bennet&Willis,2001).TwoLycopodium of the Dunes plus Grasslands plus Wetlands pollen

spore tablets were added before treatment. We sum (∑D+G+W). Finally, percentages of each

tried to reach a minimum 300 pollen grain sum pollen type grouped in Exotics are based on the

without counting Wetlands and Exotic taxa, and, in total pollen sum (∑D+G+W+E+Extra-regional).

addition, due to the overrepresentation of Poaceae Results are presented as pollen percentages and

in grasslands samples, 100 pollen grains other than include the value of pollen richness. Rarefied

Poaceae were counted. Pollen identiﬁcation was total pollen richness was performed using the

performed using the pollen reference collection minimum count size (536 pollen grains) with the

of the Paleoecology and Palynology Lab (IIMyC, Vegan package (Oksanen et al., 2017) for R (R

CONICET- Universidad Nacional de Mar del Plata) Development Core Team, 2021). Only Podocarpus

and published palynological atlases. Identified and Nothofagus t. dombeyi (extra-regional) were

pollen types were grouped into Dunes, Grasslands, excluded from the complete dataset, to calculate

Exotics, and Wetlands (Supplementary material, rariﬁed pollen richness.

Appendix 1).

Pollen vs. Landscape-scale heterogeneity analysis

Pollen - Vegetation relationships analysis

Landscape units cover (%) at a 2000 m radius

Pollen samples were classified using of each sampling site were compared using cluster

unconstrained Cluster Analysis (CONISS, TILIA analysis (CONISS, TILIA 2.6.1) results. This

2

.6.1) with square root transformation. The pollen comparison implied a two-step dimension. First,

types were organized according to habitat’s plant diﬀerent landscape unit covers were summed as: a)

species; the Wetlands group (considered as strictly low or no vegetation cover units, (<30% coverage)

local pollen), Podocarpus, and Nothofagus t. b) moderate to high vegetation cover units (>30%

dombeyi, considered an extra-regional type, were coverage), and c) exotic forest cover. Second,

excluded from this analysis.

detailed landscape units were graphed for each

In the pollen diagram, some genera or species sample site to distinguish the main landscape unit

were grouped into family categories because some contributor to pollen assemblages’.

pollen types do not enable the pollen identiﬁcation

Ordination analysis was performed to disentangle

to a lower taxonomic level (see Supplementary andquantifythemainpollentypesinﬂuencingpollen

material, Appendix 1). Non-native species of sample variance. To determine which ordination

Argentinean ﬂora were considered Exotics (Fig. method is the most appropriate for the data analysis,

3

and Supplementary material, Appendix 1) a Detrended Correspondence Analysis - DCA was

according to Cabrera & Zardini (1953), Zuloaga performed. According to Legendre & Birks (2012),

et al. (2019), and Instituto de Botánica Darwinion linear ordination methods work best for short

(2020). In Appendix 1 of Supplementary material gradients (<1.5 SD units) and unimodal methods for

the diﬀerent species that have been reported for long gradients (>3 SD units), while both methods

512

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

513

Bol. Soc. Argent. Bot. 59 (4) 2024

can be used on intermediate gradients. In this paper, list of maximum plant richness values for every

we used Principal Component Analysis (PCA) landscape unit type in Table 1). Then, we calculate

because the length of the ﬁrst DCAaxis is 1.67 units the estimated landscape plant richness at the site (푖)

of standard (SD) (closer to 1.5) therefore suggests as the sum of the maximum plant richness of every

unimodal data.

landscape unit type (푎), multiplied by its individual

cover at 1000 and 2000 m radii from each sampling

Pollen richness versus local and landscape plant site (푖).

species richness modeling

A linear regression model was applied between

A linear regression model was applied between the values of the LPRI obtained at 1000 m and 2000

local plant species richness vs. local plant species m vs. Pollen richness for each site, total as well as

richness transformed into pollen types (PSRPT), to for anemophilous and entomophilous taxa.

evaluate the eﬀect of taxonomic smoothing. Local

Finally, a linear regression model was applied

plant species richness was calculated considering between the values of axis 1 of the PCA obtained

all the diﬀerent species registered at every plot according to the pollen assemblages of sediment

included at a 20 m radius of each sampling site. samples vs. pollen richness values, as a measure

PSRPT was calculated by smoothing the plant to relate pollen-landscape unit composition with

data to pollen equivalents (plant species were sediment pollen richness values.

assigned to pollen taxa as shown in Appendix 1 of

Supplementary material).

Different linear regression models were

performed between Rareﬁed total pollen richness

Table 1. Maximum plant richness registered by

landscape unit. 5x5 m plots. Note: Atlantic Ocean,

beach, back shore and buildings and roads did not

registered plant cover so they were not considered

for this table.

a

(

henceforth pollen richness), as well as for

anemophilous and entomophilous taxa, vs. PSRPT

from plant surveys performed at 0, 10, and 20 m

radii, to evaluate possible taphonomic attributes

Maximum

Landscape units richness (spp.

number)

Vegetation

surveys per

Landscape unit

a

(

transport and deposition, and preservation) and

pollen production that aﬀect PSRPT and pollen

richness relationship.

To extend the analysis of pollen contribution

and its relationship with plant richness from local

Interdune slacks

Semiﬁxed dunes

Mobile dunes

Fixed dunes

Exotic forest

Lagoon

35

25

18

37

23

28

89

35

19

35

18

21

(0-20m) to landscape scale (0-1000 m and 0-2000

m - extra-local pollen source areas), we calculate a

Landscape plant richness Index (LPRI) following

this equation:

reSultS

RLandsc(푖): Estimated landscape plant richness at site i

푠: Number of landscape units

Pollen - Vegetation relationships analysis

Cluster Analysis of pollen samples showed

two large pollen assemblages, Group 1 and Group

푟̅푉푎: Maximum plant richness calculated for

landscape unit type a at local scale

퐶표푉푖푥푚: Ratio cover (from 0 to 1) of landscape unit

type at x meters radii from site i

푎: Semiﬁxed dunes / Fixed dunes / Mobile dunes /

Lagoon / Exotic forest / Interdune slacks

2

(Fig. 3). Group 1 presented high percentages

of Pinaceae (16.2-78.7%) and Asteraceae

subf. Asteroideae Ambrosia type (14.2-32.6%,

henceforth Asteroideae Ambrosia type), as well

as the lowest to intermediate values for Poaceae

We ﬁrst characterize the maximum plant richness (18.6-50.2%). Also, this group showed the highest

of a landscape unit type (푎) as the number of richness of exotic pollen types (10 of the 14 exotic

diﬀerent species registered considering the total pollen types registered for the complete pollen

plots surveyed for landscape unit type (푎) (see the dataset).

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C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

Group 2 was mainly characterized by high variation of the pollen data. It was explained

percentages of Poaceae (35.9-74.6%) and Adesmia mainly by the variables Pinaceae, followed by

(

0.3-23.1%). This group was further subdivided Asteroideae Ambrosia type (positive values),

into two groups (A and B). Subgroup A was Adesmia, and Poaceae (negative values). Group 1

also characterized by the highest presence of sites presented mainly positive PC1 values. The

Hydrocotyle (0.5-28.8%) and Orobanchaceae positive values also correspond to the sites with

(Agalinis/Castilleja) (0.3-10.2%). Subgroup B, in the highest percentage of exotic forest cover,

addition to Poaceae (37.3-74.6%), is characterized as opposed (negative values) to the sites with

by Pinaceae (1.1-24.6%) and presented the greatest the highest percentages of units with low or no

variety of pollen types. The sites FQ3 to FQ5, vegetation cover (Fig. 4).

recorded intermediate values of Pinaceae (13.8

PC2 axis (Fig. 5, eigenvalue 0.12) explained

-

24.6%) and high values of Poaceae. Despite 12% of variance and it was related with

the presence of species like Juncus acutus L., Poaceae (positive values), and Hydrocotyle and

Ceratophyllum sp., Triglochin striata Ruiz & Pav. Orobanchaceae (Agalinis/Castilleja, negative

in vegetation surveys (Appendix 1), no or lower values). Subgroup 2A sites presented mainly

than expected pollen grains were recorded in the negative PC2 values meanwhile Subgroup 2B

surface samples.

sites presented mainly positive PC2 values.

Pollen vs. Landscape-scale heterogeneity Pollen richness versus local and landscape plant

analysis

To obtain a general picture of pollen

species richness modeling

When we graphed Plant species richness vs.

percentages and landscape vegetation cover plant species richness transformed into pollen

relationship, we showed the information of types (PSRPT) (Fig. 6A), we observed strong

landscape units at the stacked column graphs correspondencebetweentherawdata.Theanalysis

summed as three main cover types. Units with of taxonomic smoothing (Plant species richness

low or no vegetation cover (buildings and roads, vs. PSRPT regression, see Fig. 6B) showed a

2

Atlantic Ocean, beach, back shore, and mobile positive relation (r = 0.57, p-value=1.067e-05)

dunes) vs. moderate to high native vegetation suggesting that the estimation of plant species

cover (semifixed and fixed dunes, interdune richness by pollen richness may be possible at the

slacks and lagoons), and exotic forest (Fig. 4). Coastal Dune Systems (CDS). The slope estimate

When comparing the results of Cluster Analysis (β=0.38) is considerably lower than 1, thus pollen

to the diﬀerent landscape composition at every richness values represent only ca. 40% of actual

site, for a 2000 m radius (Fig. 4A-B), Group 1 plant richness.

registered the highest percentages of exotic forest

16.77-25.52%); and high values of low or no richness (total, anemophilous, and entomophilous)

vegetation cover units (11.36-51.08%) and ﬁxed vs. PSRPT (5, 10, and 20 m) and LPRI -Landscape

dunes (17.27-45.95%). plant richness Index- (1000 m and 2000 m) are

The regressions parameters between pollen

(

Landscape units with low or no vegetation shown in Table 2. The r-squared estimate from

cover were highly represented in Subgroup 2A pollen richness vs. PSRPT regression models was

(48.12-69.76%, including mobile dunes coverage positive and signiﬁcant at 20 m, as well as at the

between 10.40-42.73%, Fig. 4A-B) with almost extra-local scale (pollen richness versus LPRI).

no exotic forest cover. Exotic forest cover was This pattern suggests that these models ﬁt best

below 23.75% at subgroup 2B (Fig. 4A-B) with the assuming a Relevant Pollen Richness Source Area

highest values at FQ3, FQ4, FQ5, and FQ23. There (RPRSA) ca. 1000 m (extra-local scale). Even

was also a mixture of landscape units with a greater though 64% of total plant species recorded in

representation of interdune slacks in some sites and this study present entomophilous pollination,

ﬁxed dunes in others.

entomophilous pollen richness versus plant

The PCA (Fig. 5) showed two large groups richness relationship was not signiﬁcant at any

mostly consistent with Cluster Analysis. PC1 of the scales, while anemophilous pollen richness

axis (eigenvalue 0.29) described 29% of the versus plant richness relationship presented

515

Bol. Soc. Argent. Bot. 59 (4) 2024

Fig. 4. Stacked columns graph of landscape units cover (%) at 2000 m radius, ordered according to CONISS

analysis. 4B. Low or no vegetation cover units (<30%); moderate to high vegetation cover units (>30%).

516

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

Fig. 5. Principal Component Analysis biplot, with squared transformation, for the diﬀerent sampling sites

according to the pollen types, and the loadings (eigenvectors), on the ﬁrst two principal component axes.

The main groups separated by CONISS (Fig. 4) are delimited in black circles (1 and 2), and the subgroups

2A and 2B are delimited in pink and orange respectively. Orobanchaceae = Orobanchaceae (Agalinis/

Castilleja). For visual purposes, we only show the 10 variables that contribute the most (>3%) to PC1 or PC2.

Table 2. Linear regression results between PSRPT, and LPRI vs. Pollen richness. Abbreviations= PSRPT: plant

species richness transformed into pollen types; LPRI: landscape plant richness Index. *Signiﬁcance 0.05.

Scale of analysis– Pollen

Variables evaluated

Adjusted R-squared

p-value

source area scale

m - Local

10 m - Local

5

-0,04

-0,03

0,16

-0,01

-0,04

-0,008

-0,02

0,01

0,39

0,27

0,07

0,18

0,28

0,04

0,14

0,8

0,54

PSRPT vs. Pollen richness

20 m - Local

0,03*

0,42

5

m - Local

0 m - Local

PSRPT vs. Pollen richness

Entomophilous

1

2

0,78

0,37

5

m - Local

10 m - Local

0 m - Local

0,52

PSRPT vs. Pollen richness Anemophilous

0,28

2

0,002*

0,006*

0,11

LPRI vs. Pollen richness

1

000 m (landscape)

extra-local

LPRI vs. Pollen richness Entomophilous

LPRI vs. Pollen richness Anemophilous

LPRI vs. Pollen richness

-

0,02*

0,005*

0,17

2

000 m (landscape)

extra-local

LPRI vs. Pollen richness Entomophilous

LPRI vs. Pollen richness Anemophilous

-

0,04*

517

Bol. Soc. Argent. Bot. 59 (4) 2024

Fig. 6. The loss of plant species richness due to taxonomic smoothing to plant genus or family levels and

pollen types equivalents. A. Area graph that shows the magnitude of the diﬀerences according to Plant

spp. richness vs. Plant spp. richness (in pollen types-PSRPT) categories for each pollen sample. B. Linear

regression graph between Plant spp. richness vs. PSRPT. β estimate is shown at the modelled regression

equation.

518

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

positive and signiﬁcant adjusted r2 values at the presented higher pollen richness values than pure

2

0 m and extra-local scales.

native environments (those with high values of

Comparison between pollen richness values Adesmia and Poaceae pollen percentages), see Fig.

and diﬀerent pollen assemblages suggested that 5 and Table 3. Indeed, the average pollen richness

sites with the presence of exotic forest (those with of the sites in Group 1 (Fig. 5) was 33.4, while for

high values of Pinaceae and Asteroideae Ambrosia those in the middle zone (subgroup 2B) it was 29

type pollen percentages) within a 2000 m radius and for those on the right (subgroup 2A) it was 28.

Table 3. Linear regression results between PC1 Axis vs. Pollen richness and the pollen variables that

most contribute to PC1, with squared transformation. * Signiﬁcance0.05.

Variables evaluated

Adjusted R-squared

Adjusted R-squared p-value

b

b p-value

PC1 Axis vs. Pollen richness

PC1 Axis vs. % Pinaceae

PC1 Axis vs. % Ambrosia

PC1 Axis vs. % Adesmia

PC1 Axis vs. % Poaceae

0.07

0.8

0.11

1.643e-09*

5.057e-06*

0.001*

0.59

0.71

0.4

0.11

1.64e-09*

5.06e-06*

0.001*

0.6

0.34

0.18

-0.29

-0.2

0.02*

diScuSSion

vegetation relationships at a local scale or compared

qualitatively diﬀerent vegetation units. This study

presents the ﬁrst quantitative analysis of pollen

Pollen - Vegetation relationships analysis

Do surface sediment pollen assemblages and vegetation relationship in terms of spatial

reﬂect landscape heterogeneity at the CDS?: The heterogeneity at an extra-local scale (2000 m radius

challenge of this study was to assess whether pollen from a sampling site). The pollen-vegetation model

assemblages reflect the heterogeneity of plant presented here shows that pollen assemblages

communities within the Coastal Dune Systems may discern between forested and non-forested

(

CDS) area. We found that pollen assemblages landscapes. Furthermore, mobile dunes vs. ﬁxed

were strongly inﬂuenced by landscape unit cover, dunes+semi-ﬁxed dunes+interdune slacks could

beyond the local scale, and reﬂected landscape be separated as well. These results about the

heterogeneity. This is mainly reﬂected in the pollen attributes of the pollen record may be compared

composition of sampling sites with predominantly with other coastal dune systems to evaluate the

exotic forest, mobile dunes or mixed landscape effect of different vegetation compositions on

units.

Several studies carried out in coastal dunes

pollen representation.

We found that pollen assemblages from sites of

ecosystems in Argentina, Uruguay and in small the CDS surrounded by exotic forests showed high

lake in the northeastern United States, have also values of Pinaceae and Asteroideae Ambrosia type

found correspondence between plant communities (e.g. Group 1: FQ6-FQ8, Fig. 3 and Fig. 4). The

and environments, and confirmed that pollen relevant values of Pinaceae reported in this study

spectra reﬂect the spatial heterogeneity of diﬀerent had already been highlighted in previous steppe

plant communities (Stutz & Prieto, 2003; Fontana, environments (Li et al., 2022), considering also

2

005a; Marcos & Mancini, 2012; Montserrat that e.g. Pinus spp. has a good pollen dispersal and

et al., 2012; Masciadri et al., 2013; Liu et al., is usually over-represented in pollen assemblages

022). All these studies have evaluated pollen and (Felde et al., 2015). Also, the large amount of

2

519

Bol. Soc. Argent. Bot. 59 (4) 2024

Pinaceae may be related to a) higher pollen by various adventitious species (Cabrera, 1941).

productivity than native herbs and shrubs of the This could explain the lower values of Adesmia in

CDS and b) the regional pollen source area is Group 1, both at the pollen record and plant cover.

composed mainly of landscape units with low or Hydrocotyle bonariensis is a frequent hydrophytic

no vegetation cover. Likewise, it has been possible species in sandy and humid soils that adapts

to evidence that aﬀorestation with exotic species easily to sandy soils, as do Agalinis communis

reduces the area of psammophytic grassland (Yezzi and Castilleja arvensis, adapted to coastal dunes

et al., 2018) which implies less participation of (Cabrera, 1941). The other sites -Group 2B- showed

other pollen types.

the highest values of Poaceae and Asteraceae.

The major vegetation taxa belong to Poaceae,

Sites FQ3 to FQ5 with intermediate percentages

of exotic forest landscape unit (in a 2000 m radius) Cyperaceae, and Asteraceae in this study coinciding

registered intermediate values of Pinaceae, but with those results reported by Fontana (2005a) in the

they also recorded high values of Poaceae, both southern CDS. The higher values of Poaceae could

pollen types with anemophilous dispersal. As be because this family includes several species

already stated, the composition of pollen spectra and had representatives in all landscape units.

is inﬂuenced by dominant wind patterns (Fontana, Likewise, they have a high pollen production, with

2

005a) since high wind speed could carry more an anemophilous and easily dispersed pollen grain,

pollen grains (van der Sande et al., 2021). In that favors their good representation in the pollen

these sites, landscape units with moderate to high spectra (Fernández & Grill, 2016). Stutz & Prieto

vegetation cover dominate south/southwestward (to (2003) also highlighted that Poaceae, Cyperaceae,

sampling sites).As Faro Querandí Municipal Nature and psammophytic types (like Adesmia incana,

Reserve (FQNR) presented a south/southwest Calycera crassifolia, Senega cyparissias (A. St.-

dominant wind direction, this factor would Hil. & Moq.) J.F.B. Pastore & Agust. Martinez

counteract the over-representation of Pinaceae and (Polygala type), Apiaceae -including Hydrocotyle

favored the contribution of Poaceae and other taxa. - and Asteraceae which grow on sandy soils),

On the contrary, in Group 1, the highest exotic conformed the pollen assemblages of the coastal

forest cover was located north/northwestward, dune barrier in the area of the Mar Chiquita coastal

which favored the over-representation of Pinaceae.

lagoon, adjacent to the FQNR. Also, Fontana

We found a strong Asteroideae Ambrosia type (2005a) pointed out that pollen assemblages

association with sites located in patchy grassland- differed considerably from the composition of

forested areas. This was consistent with post surrounding local vegetation. These results are

disturbance colonization of Ambrosia associated coherent with those reported in this study as

with pine forest (Yezzi et al., 2018). These high pollen assemblages from surface samples reﬂect

values of Ambrosia pollen type in the CDS have not only local vegetation, but indeed they reﬂect

already been reported in previous studies such as a wider pollen source area and sense landscape

that of Stutz & Prieto (2003) and Fontana (2005a), heterogeneity.

although these authors did not investigate the

association of Ambrosia pollen type with pine psammophytic pollen types in addition to Poaceae

forests. and Asteraceae (like Apiaceae, Caryophyllaceae,

Group 2B recorded a greater variety of

We also found that sampling sites surrounded and Calycera). These sites also had a mix of

up to 40% of landscape units with low vegetation landscape units, mainly ﬁxed/semi-ﬁxed dunes and

cover (e.g. Mobile dunes, Group 2A) had a strong interdune slacks. This situation was similar to that

inﬂuence of psammophytic species pollen types, reported by Montserrat et al. (2012), who found

like Adesmia, Hydrocotyle, and Orobanchaceae that the sectors located on these landscape units

(

Agalinis communis (Cham. & Schltdl.) D’Arcy were characterized by diverse plant associations,

and Castilleja arvensis Schltdl. & Cham.), genera and the species composition was highly diverse.

and species which have entomophilous pollen This great vegetation heterogeneity and the eﬀect

dispersal. Adesmia incana, a psammophytic species of taxonomic smoothing, would trigger the pollen

abundant in sandy dunes, can be affected by assemblages of these landscape unit overlap. It

indirect anthropogenic action, being substituted is not possible to separate at the genus or species

520

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

level of Poaceae and Asteraceae, pollen types that Sugita, 1994; Odgaard, 2018). The relation between

could be indicative of each type of environment. local and extra-local plant and pollen richness

These diﬃculties have been previously reported in reported in our study may reﬂect vegetation structure

southern CDS (Fontana, 2005a) and dune deserts in and diversity at these scales, rather than at the

north China (Li et al., 2022).

regional level, as previously published by Goring

et al. (2013), Matthias and Giesecke (2014) and

Matthias et al. (2015). Odgaard (2018) highlighted

Pollen, plant and, landscape richness analysis

The relationship between plant richness and that modern pollen assemblages can better reﬂect α,

pollen richness is driven both by taxonomic γ, or ε- diversity according to the size of the lakes

smoothing, pollen production, and taphonomic or basin where they have been deposited. Sugita

constraints (pollen dispersal and preservation). The (2007a) indicates small lakes (up to 350 m radius,

eﬀect of taxonomic smoothing is likely due to the like those of this study) are more suitable to capture

abundance of plant species producing pollen grains local vegetation spatial heterogeneity (ca. 700 m)

often only identiﬁable to genus, and sometimes to by the 30-45% of the total pollen loading deposited

family, like Poaceae, Asteraceae and Cyperaceae. in the basin. However, pollen richness calculation

In this study, Panicum racemosum, Poa lanuginosa, is based on the pollen counts of the complete pollen

and Polypogon monspeliensis (L.) Desf. were loading deposited in a basin. That includes those

registered in the vegetation surveys, but at the pollen pollen types from the local scale (the Relevant

level they could only be identiﬁed as Poaceae. Also, Pollen Source Area- RPSA- sensu Sugita, 2007a)

Eleocharis maculosa (Vahl) Roem. & Schult., and those beyond the RPSA from extra-local. Thus,

Isolepis spp., and Schoenoplectus americanus ﬁrstly, the fact that our models present signiﬁcant

(Pers.) Volkart ex Schinz & R. Keller were recorded adjusted-r2 values for local scales pollen-plant

in the vegetation but their pollen grains could only richness relationship coincides with those results

be determined at the family level: Cyperaceae. The previously published by Sugita (2007a). Secondly,

ﬁdelity of pollen richness relative to plant richness our results suggest that the RPRSA ﬁts with extra-

may be aﬀected by taxonomic smoothing since the local scale pollen-plant richness relationships. This

taxonomy in the plant dataset is not equivalent to inference coincides with previously published

pollen taxonomy, but this eﬀect may be recoverable semiquantitative-qualitative pollen-vegetation

(Goring et al., 2013). Thus, taxonomic smoothing relationships studies for the CDS (e.g. Stutz &

of CDS plant richness could be overcome by the use Prieto, 2003;Latorreetal., 2010).ThewiderRPRSA

of the statistical models presented in this paper and than expected RPSA may be related to extra-local

adding diﬀerent paleobotany proxies to the classic pollen inputs linked to a) anemophilous species

vegetation-palynological based reconstruction (e.g. Poaceae and Amaranthaceae/Chenopodiaceae

when studying Holocene records.

species) growing in open landscapes as semiﬁxed

As we discussed in the previous section (“Pollen- and mobile dunes (Latorre et al., 2010; Felde et

Vegetation relationships analysis”) that pollen al., 2015; Adeleye et al., 2020) and b) species

assemblages reﬂect landscape heterogeneity, the with high pollen productivity (Latorre, 1999;

Relevant Pollen Richness Source Area (RPRSA) Odgaard, 1999; Weng et al., 2006) growing in

for small lagoons of the CDS correlate with extra- forest patches (e.g. Ambrosia tenuifolia, Pinus spp.)

local scale vegetation heterogeneity. Even though of the CDS. Weng et al. (2006) and Goring et al.

the regression models between plant and pollen (2013) have also highlighted that anemophilous

richness were signiﬁcant at the local scale (20 pollen types are usually the main component of

m, Table 2), the goodness of ﬁt of these models pollen assemblages and drive pollen richness

increase and stabilize up to the extra-local scale patterns. Some entomophilous and hydrophytic-

(1000 and 2000 m, Table 2). Recent studies have anemophilous species are also important local

shown that modern pollen assemblages can reﬂect scale components of pollen richness. Between

plant richness at diﬀerent scales (Birks et al., 2016), entomophilous species we found Asteraceae

however, the relationship is not direct as pollen (Tessaria type, Baccharis type, Senecio type, and

richness reflects pollen evenness, productivity, other Asteroideae), Hydrocotyle spp., and Adesmia

and dispersal across the landscape (Prentice, 1985; incana which pollen grains can be transported over

521

Bol. Soc. Argent. Bot. 59 (4) 2024

a few meters (Wang et al., 2022), they usually et al. (2020) for Patagonian forest-steppe ecotone

produce low quantity of pollen grains (Väli et and for Central Europe by Abraham et al. (2020).

al., 2022) and are under-represented in pollen Birks & Line (1992) and Matthias et al. (2015) have

assemblages (Mourelle et al., 2017). Hydrophytic- discussed the relationship between disturbance

anemophilous species growing at interdune slacks frequency and pollen richness. They stay that a

(Cyperaceae species and Typha spp.) produce high disturbed environment is related to higher pollen

quantities of pollen which are highly dispersed (Xu richness than a stable environment. Disturbances

et al., 2012; Frazer et al., 2020). This fact explains promote the availability of new niches and thus

the high values of these pollen types reported in the establishment of new species. If disturbance

most sites of the CDS even though these species frequency is high (low), vegetation communities

may be absent in some interdune slacks.

are expected to be dominated by r-strategy plants

Pollen richness is also a function of landscape (k-strategy plants). Intermediate disturbance

structure, openness and plant diversity within frequencies favor the coexistence of both r and

the Relevant Pollen Source Area (RPSA), k- strategy plants, thus showing the highest

pollination syndromes and dispersal, and other species richness values (“intermediate disturbance

taphonomic processes (Birks et al., 2016). Within hypothesis”, Wittaker et al., 2001). The higher

these taphonomic processes, we found that pollen pollen richness values linked to sampling sites with

preservation could be diﬀerent between taxa and values of exotic forest up to 20% may be associated

could be a source of uncertainty (Goring et al., with coastal dune environments characterized

2

013; Grindean et al., 2019; Li et al., 2022; Wang by intermediate disturbance frequencies regime.

et al., 2022). Regarding taphonomic processes The establishment of small patches of forest

triggering pollen-vegetation representation, we communities may diminish the frequency

found that Juncaceae and Juncaginaceae pollen of vegetation burial because of they partially

types as those with minor preservation potential prevent dune movements and provides a mosaic

in the CDS. As noted in the results section, in the of heliophilous and shade-tolerant environments.

vegetation surveys, species of these families were Also, anthropogenic fires, vegetation clearings

recorded although they were absent from the pollen linked to oﬀ-road tourism, and horse sheltering

record in the ﬁrst case, or with low representation usually increase near forested areas, and thus these

in the second. These and other taxa of hydrophilic types of disturbance frequency. Conversely, sites

pollen present very thin exines that hinders their with the lowest pollen richness were found in more

preservation, and also tends to disintegrate during pristine areas with a high presence of landscape

acetolysis (Moore et al., 1991), therefore, causes units with low or no vegetation cover, mainly

their low representation in the pollen records mobile dunes, and with a low presence of the exotic

(Fontana, 2005a; Erdtman, 1952; Chenlo, 2014).

forest. This low plant richness (and consequently

The higher pollen richness values recorded at low pollen richness) may be due to the strong

sampling sites with high values of exotic forest (up stress conditions that occur in active dunes, such

to 20% coverage within a 2000 m radius) is one of as high wind intensity, substrate instability, saline

the most outstanding results we reported for the spray, and low organic matter content, among

CDS. All these sites presented a forest matrix mixed others, which cause that few species can adapt to

with open grasslands areas with diverse herbaceous these conditions (Kumler, 1997; Fontana, 2005a;

and shrub species, which were characterized by Montserrat et al., 2012).

more stable substrate and sheltered from wind

desiccation effects. These factors favor higher

organic matter content accumulation and less concluSionS

frequency of disturbances related to burial eﬀects

produced by sand over vegetation, which may also

favor greater species richness (Montserrat et al., type cover, patchiness, and location with respect

012). The increase in pollen richness associated to dominant wind direction), pollen production,

Landscape heterogeneity (in terms of vegetation

2

to the development of forest patches with open taphonomic constraints and taxonomic smoothing

steppe environments was also reported by Sottile determined pollen assemblages from small lagoons

522

C. Vásquez et al. - Pollen representation of vegetation and plant richness of pampean coastal dunes

in Pampean CDS. Pollen assemblages are good 2021 -11220200100204CO), SS. We also thank to

proxy records to reconstruct past landscape the park rangers of the Faro Querandí Municipal

heterogeneity at an extra-local scale (ca. 2000 m) Nature Reserve, for their help during the ﬁeld work

of coastal grasslands of southern South America. and two anonymous reviewers whose comments

As other grassland ecosystems (dominated by contributed improving a previous version of this

Poaceae, Asteraceae and Cyperaceae), pollen manuscript.

richness represents less than 50% of real plant

richness. However, we reported a signiﬁcant relation

between plant and pollen richness as a promising Primary data availability

proxy record encouraging future landscape scale

paleoecological reconstructions. Our results suggest

All plant species recorded in this work

that anemophilous pollen types play a major role are available at https://www.gbif.org/es/

in driving pollen richness, and that a landscape dataset/36fc65f3-ba8d-4c06-917e-6ffe62b63577

composed of coastal dunes and small forest patches and pollen information is deposited in Repositorio

provideshighervaluesofpollenrichnessthanpristine Institucional CONICET (https://ri.conicet.gov.ar/).

coastal dune patches. Even though we detected few

entomophilous species that frequently contribute to

pollen richness, we encourage the consideration of bibliograPhy

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anemophilous or entomophilous) to draw a better ABRAHAM, V., J. ROLEČEK, O. VILD, E.

picture of the paleoenvironmental changes. Future

studies may explore other techniques that avoid the

use of strong acid-base solutions that may aﬀect

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such as Juncaceae or Juncaginaceae. We presented

JAMRICHOVÁ … & P. KUNEŠ. 2020. Spatial

scaling of pollen-based alpha and beta diversity

within forest and open landscapes of Central Europe.

bioRxiv: 2020.08.18.255737.

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HABERLE … & J. STEVENSON. 2020. Long-

term drivers of vegetation turnover in Southern

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