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Quaternary Science Reviews

journal homepage: www.elsevier.com/locate/quascirev

Riparian woody vegetation history in the campos region, southeastern

South America, during two time windows: late Pleistocene and late
Holocene
Dominique Mourelle a, *, Aldo R. Prieto b, Felipe García-Rodríguez a
a
Centro Universitario Regional Este, CURE-Rocha, Uruguay
b
IIMyC, CONICET-Universidad Nacional de Mar del Plata, Facultad de Ciencias Exactas y Naturales, Laboratorio de Paleoecología y Palinología, Funes 3250,
7600, Mar del Plata, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: A detailed palynological record from Laguna Formosa (northeastern campos region, 31
S; 54
W) docu-
Received 13 March 2017 ments the dynamic balance between grasslands and riparian forests during the late Pleistocene (14,570
Received in revised form to 13,500 cal yr BP) and late Holocene (3280 cal yr BP to the present). Modern pollen-vegetation re-
25 April 2017
lationships and the woody pollen dispersal capacity analyses were used to improve the vegetation
Accepted 26 April 2017
reconstruction. Grasslands were regionally dominant throughout the record. However, at 14,570 cal yr BP
hydrophilous taxa reflect the development of riparian hydrophilous shrublands along freshwater bodies,
promoting the fixation of the riverbanks, maintaining shallow, calm and clear water conditions under a
Keywords:
Pleistocene
relatively wet and not so cool climate. This is the first evidence of woody riparian vegetation develop-
Holocene ment along freshwater bodies for the lowlands of the northern campos during the late glacial period. At
Palynology 3280 cal yr BP riparian forests consisted of both hydrophilous and mesophilous woody taxa. Since
Riparian forests 2270 cal yr BP woody vegetation gradually increased, accompanied by the incorporation of other taxa by
Rio de la Plata grasslands 940 cal yr BP, and achieving a composition similar to that of the contemporary time at ca. 540 cal yr BP.
Campos The increased woody vegetation since ca. 2270 cal yr BP, and the more frequent and intense flooding
Uruguay events between 1800 and 1200 cal yr BP, could be related to higher precipitation over La Plata Drainage
Brazil
Basin, related with the high ENSO amplitude. In addition, pollen from taxa that currently no longer
Araucaria
develops in the study area suggests connections between southern Brazil and Uruguay, and between the
campos and the Chaco phytogeographic province.
© 2017 Elsevier Ltd. All rights reserved.

1. Introduction to certain topographic features that provide relatively high water

The campos region in Uruguay and southern Rio Grande do Sul
(Brazil) and the pampas region in the eastern Argentina integrate
the Río de la Plata grasslands (RPG) (700,000 km2), the main
complex of grassland ecosystem in South America (Soriano, 1991)
(Fig. 1). This region displays a conspicuous biodiversity, with ca.
3000 species of vascular plants, including more than 550 different
grass species from very diverse genus. The particular climatic
conditions of the region allow the coexistence of many species with
C3 and C4 metabolisms (Bilenca and Min ~ arro, 2004).
In the campos region, trees and shrubs are frequent but limited
* Corresponding author.
E-mail addresses: domodica@gmail.com (D. Mourelle), aprieto@mdp.edu.ar
(A.R. Prieto), felipegr@fcien.edu.uy (F. García-Rodríguez).
availability or wind protection, such as corridors along stream and
river basins and rocky slopes. The woody flora of the Uruguayan
campos consists of >300 species (Brussa and Grela, 2007; Haretche
et al., 2012), therefore some authors considered that Uruguay
should not be fully included in the RPG (Chebataroff, 1960; Grela,
2004). According to Grela (2004) the relatively narrow strip on
the eastern part of the Río Uruguay should be considered a tran-
sition boundary between the Chaco and the Paranaense phyto-
geographical provinces (Fig. 1). This assumption is supported on
woody plant distributions supported by both the penetration of
tropical vegetation along this river and the development of species
from Chaco phytogeographic province in the adjacent plains.
Some paleoecological studies in the campos region of southern
Rio Grande do Sul and Uruguay have been carried out in the last
decade. Palynomorph and phytolith data from the Rio Grande do

http://dx.doi.org/10.1016/j.quascirev.2017.04.024
0277-3791/© 2017 Elsevier Ltd. All rights reserved.
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29 15

~o central gaúcha and modern Araucaria populations; c)

Fig. 1. a) Location map of campos region with sites mentioned in text; b) NE Uruguay and SW Rio Grande do Sul, depressa
upper reaches region of the Río Negro (URRN) showing the location of Laguna Formosa; d) surface pollen samples and LFOR1 core location.

Sul lowlands (S~ao Francisco de Assis: Behling et al., 2005) and
Uruguay (Paso Barranca-India Muerta: Iriarte, 2006; Mourelle et al.,
2015a; Laguna Negra: García-Rodríguez et al., 2010; Arroyo Solís
rez, 2011) suggest
Grande: Mourelle et al., 2015b; and Pay Paso: Sua
that grasslands were the predominant physiognomy during the late
Pleistocene and early-mid Holocene (Fig. 1). Riparian (gallery)
forests began to develop along streams and rivers at ca. 5170 cal yr
BP in Rio Grande do Sul lowland campos, and they expanded from
ca. 1550 cal yr BP (Behling et al., 2005). For the Uruguayan campos,
grasslands were dominated by C3 grasses during late Pleistocene,
and were replaced by C4 grasses during the Holocene (Iriarte, 2006)
(Fig. 1). Pollen analysis suggests the beginning of the expansion of
16 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
forest along the riverbanks at the northwestern Uruguayan campos
(30
S; 57
300 W) from ca. 12,400 cal yr BP (Sua rez, 2011) (Fig. 1).
However, such paleoecological interpretation should be considered
with caution because the palynological information was derived
from three archaeological sites, where most of the samples exhibit
not only very low total pollen sums (<50 pollen grains), but also up
to 40% of the pollen grains were deteriorated. In the southeastern
part of this region the development of riparian forest patches in
river basins since ca. 8000 cal yr BP has been suggested by other
palynological studies (Iriarte, 2006; Mourelle et al., 2015a,b) (Fig. 1).
The dominance of grasslands vegetation in the Rio Grande do
Sul southern lowlands during the late Pleistocene and early-mid
Holocene is associated to cold and relatively dry, and warm and
dry climatic conditions, respectively (Behling et al., 2005) (Fig. 1).
The authors refer to a possible sedimentary hiatus between 220 cm
and 195 cm core depth, as suggested by the radiocarbon dates and
the sediment stratigraphy, which would correspond to the early
and/or mid-Holocene. However, since pollen preservation was very
poor, there is no reliable palynological information for this interval
(Behling et al., 2005). Despite these facts, the pollen sequence was
interpreted as a controversial continuous formation. After ca.
5170 cal yr BP, wetter climatic conditions were inferred by the
initial expansion of riparian forests in the Rio Grande do Sul low-
lands (Behling et al., 2005). For the Uruguayan campos, the C3
grasses dominance during late Pleistocene suggests drier and
cooler climatic conditions (Iriarte, 2006). During the early and mid-
Holocene, warmer and more humid and drier conditions, respec-
tively, have been suggested by Iriarte (2006), followed by humid
conditions in the late Holocene (Iriarte, 2006; Mourelle et al.,
2015a). This climatic variability promoted changes in the grass-
land vegetation at local scales, related to changes in inland fresh-
water availability (Mourelle et al., 2015a) and coastal change
subject to relative sea-level change (García-Rodríguez et al., 2010;
Mourelle et al., 2015b).
Palynological studies of the southern Brazilian Eastern Plateau
(highlands) indicated: (1) a predominantly cold and dry climate at
the end of the late Pleistocene, with grassland vegetation and forest
taxa restricted to refuges; (2) increased temperature and humidity
in the early Holocene, with the onset of the forest expansion from
refuges towards grasslands; (3) a dry phase in the mid-Holocene
with decreased vegetation cover; and (4) increased temperature
and humidity from the late Holocene to the present, supporting the
development of forest ecosystems (expansion of Araucaria in the
highland and tropical rainforest along the coastal slopes of moun-
tain chains) (Roth and Lorscheitter, 1993; Leonhardt and
Lorscheitter, 2010; Jeske-Pieruschka and Behling, 2012; Scherer
and Lorscheitter, 2014). However, the temperature increase during
this phase probably reduced the reproductive capacity of the
Araucaria forest taxa, thus resulting in a lower rate of expansion in
the grassland (Scherer and Lorscheitter, 2014; Spalding and
Lorscheitter, 2015). In fact, no events of major vegetation change
were observed during early Holocene within this region. Instead,
vegetation changes were predominantly inferred for the last
4000 yr (Rodrigues et al., 2016). Some authors agree with this
general interpretation but still indicate a dry climate at the begin-
ning of the Holocene (Behling et al., 2001, 2004).
There is a limited knowledge about the riparian woody vege-
tation history in the campos region, except for some mention of
riparian forest patches developed during the Holocene in the
southeastern Uruguay (Iriarte, 2006; Mourelle et al., 2015a,b) and
the little information for Rio Grande do Sul lowlands (Behling et al.,
2005). The upper reaches region of the Río Negro (URRN) (Fig. 1)
represents a key area to investigate the late Quaternary woody
vegetation development and climate dynamics in the RPG, since it
is a very important wetland area in the lowlands of northern
campos, where riparian forests are both well developed and sur-
rounded by dominant grasslands and practically no anthropogenic
impacts have disturbed the environment. Our goal is infer the
woody vegetation development and the environmental history
during two time windows, from the palynological record of Laguna
Formosa. We used modern pollen-vegetation relationships and the
pollen dispersal capacity of the most frequent woody taxa to
improve the vegetation reconstruction. In particular we aim to
integrate our results to other palynological and fossil vertebrate
records in order to discuss the possible phytogeographical con-
nections overall the campos region of RPG.
2. Study area
2.1. Environmental setting
Laguna Formosa is a shallow lake that developed from an
abandoned meander situated in the URRN (Fig. 1). The Río Negro is
the most important lotic system in Uruguay, that rises in Sierra de
Santa Tecla highland (southern Brazil) and flows westwards across
the entire country to the Río Uruguay (Fig. 1) (Praderi and Vivo,
1969). Despite the construction of Rinco n del Bonete and Baygorria
hydroelectric dams changed many sectors of the basin, the URRN
remains unchanged (Praderi and Vivo, 1969). In URRN there are
many abandoned meanders that have resulted from the Río Negro
stream channel cut off, such as Laguna Formosa, one of the most
extensive of the URRN with a surface area of ca. 28.5 ha and a
maximum depth of 1.5 m (Fig. 1).
According to Leo  n (1991) the study area corresponds to northern
campos region of the RPG (Fig. 1), characterized by grasses from
Andropogoneae and Paniceae tribes (e.g. Andropogon, Schizachy-
rium, Axonopus and Paspalum). In addition, riparian (gallery) forests
locally develop, in narrow strips along the margin of the water
streams and surrounding swampy areas and freshwater bodies
(Fig. 2). The main taxa are Salix chilensis, Phyllanthus sellowianus,
Cephalanthus glabratus, Sebastiania commersoniana, Erytrina crista-
galli, Acanthosyris spinescens, Scutia buxifolia, Myrsine laetevirens,
and many Myrtaceae (Mourelle and Prieto, 2012, 2016). Freshwater
streams and lakes are also surrounded by hydrophilous shrubs,
dominated by Phyllanthus sellowianus, Cephalanthus glabratus and
Sebastiania schottiana. Emergent aquatic plants rooted onto the
littoral shallowest areas are frequent, and consists of either a single
species, such as Ludwigia peploides or Polygonum punctatum, or
many taxa, such as Eichhornia spp., Pontederia sp., and Nymphoides
indica. Free floating Azolla filiculoides and Pistia stratiotes also
develop freely on the surface. The relief of the URRN is gently-
rolling, interrupted by some rock outcrops; there is a rich fluvial
network where streams and rivers are commonly surrounded by
the riparian forests (Leo  n, 1991). This region is called as the
depressa ~o central gaúcha, and operates as a corridor of floristic
connection between southern Brazil and northeastern Uruguay
(Chebataroff, 1960; Brussa and Grela, 2007) (Fig. 1). As a conse-
quence, riparian forests in northeastern Uruguay contain some taxa
that only develop in this area (e.g. Mimosa spp., Xylosma spp. and
Citronella spp.) and achieve higher species richness than those of
the forests from other regions of Uruguay (Brussa and Grela, 2007).
Wet meadow herbaceous vegetation develops between the ri-
parian forests and the water bodies. It consists of grasses, emergent
aquatic plants such as Cyperaceae, Polygonum and Alternanthera,
and other small herbs frequently from Asteraceae and Boraginaceae
families. In the grassland-forest boundary, bunchgrasses and
freshwater marshes are also frequent (Fig. 2).
In the grassland-forest boundary, bunchgrasses are dominated
by Panicum prionitis and Paspalum quadrifarium, and isolated trees
(e.g. Erythrina crista-galli) and shrubs (e.g. Solanum glaucophyllum,
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29 17

Fig. 2. a) Palynomorph percentage diagram of the surface soil samples from URRN region; b) schematic representation of URRN vegetation types where Laguna Formosa is located;
each pie graph represent grasslands þ bunchgrasses, Asteraceae subf. Asteroideae and riparian forest pollen proportion in each surface sample; c) pollen proportion of the most
important riparian woody taxa in surface samples.

Sesbania punicea) are also observed. In small areas with perma-
nently high groundwater, there are freshwater marshes with
aquatic vegetation (Fig. 2) (Mourelle and Prieto, 2016).
2.2. Climatic features
Present climate at URRN is temperate with seasonal mean
temperatures ranging from 12
C in winter to 23
C in summer due
to the annual cycle of insolation. The lowest measured monthly
average temperature is 5
C in June, and frosts commonly occur in
spring. Total annual precipitation varies from year to year, but mean
historical total rainfall is 1400 mm yr1 and surface mean wind
speed is about 3.5e4.5 m s1 (e.g. Pisciottano et al., 1994; Diaz et al.,
1998; DNM, 2017). Regional atmospheric circulation is under the
18 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
influence of the South Atlantic semi-permanent high-pressure cell,
responsible for not only the transport of considerable moisture over
the eastern La Plata Drainage Basin but also for the seasonal peak of
precipitation in austral summer, when the southward moisture
flow from Amazonia and convective processes forced by conti-
nental solar heating are more intense. Reinforced (weakened)
South America convergence zone (SACZ) and cold (warm) water in
the Southwestern Atlantic can reduce (increase) summer precipi-
tation in most of Uruguay (Doyle and Barros, 2002; Díaz and
Aceituno, 2003). In addition, studies of ENSO-related rainfall
anomalies reported strong influence of these events over Uruguay
during late spring-early summer and late fall-early winter (e.g.
Pisciottano et al., 1994; Diaz et al., 1998; Cazes-Boezio et al., 2003).
3. Materials and methods
3.1. Sampling and sediment analyses
In March 2011, a 163-cm-long sediment core was collected
with the aid of a piston corer from the southern border of Laguna
Formosa, at 80 m equidistant from the margins (LFOR1;
31
480 4300 S; 54
280 4100 W; 123 m a.s.l.) (Fig. 1). The core was
opened in the laboratory, described in detail and sectioned every
1-cm intervals. Subsamples were analyzed for: (1) pollen and
spores; (2) non-pollen palynomorphs (NPPs; includes algae and
cyanobacteria); and (3) organic matter content. Organic matter
content was estimated by loss on ignition (LOI) (Heiri et al.,
2001).
3.2. Palynological analysis
In order to perform an analysis of the local pollen representation
in URRN, eight surface soil samples from riparian forests, bunch-
grasses and wet meadow vegetation were collected following
Adam and Mehringer (1975) sampling strategy (Fig. 2).
Surface pollen samples were dried at 60
C for 24 h, and sub-
samples of 5 g dry sediment were used for pollen analysis,
whereas for fossil pollen and NPPs analyses samples of 2e3 cm3 of
sediment were taken at predetermined intervals along the core.
Standard palynological techniques were performed for palyno-
logical extraction using KOH, HCl, ZnCl2 for heavy liquid separa-
tion, HF and acetolysis (Faegri and Iversen, 1989). Three
Lycopodium clavatum spore tablets were added before treatment
to calculate pollen concentration (Stockmarr, 1971). Pollen and
spores were identified using atlases and published keys (Prieto
and Quattrocchio, 1993; Bauermann et al., 2013), and the mod-
ern reference pollen collection at the Laboratorio de Paleoecología
y Palinología, Universidad Nacional de Mar del Plata, Argentina.
Identification of NPPs was based on van Geel (2001), Borel et al.
(2003) and Borel (2007).
At least 300 pollen grains, excluding CYPERACEAE and aquatic taxa
due to its local over-representation, were identified and counted for
all samples. Each pollen type was calculated as the percentage of
the total pollen sum. NPPs were calculated as the percentage of the
pollen sum plus NPPs sum. Bryophytes, lycophytes and mon-
ilophytes were calculated as percentages of the pollen sum plus
spores sum. Vascular plant nomenclature follows the nomenclature
system of the Institute of Botany Darwinion of Argentina (http://
www.darwin.edu.ar) and of the Missouri Botanical Garden
(http://www.tropicos.org/). Since there are significant differences
between the concepts of “plant taxa” and “palynomorphological
types” and in order to provide the necessary clarity on the
nomenclature, names of pollen and spores were represented with
SMALL CAPITALS and refrain from using italics (de Klerk and Joosten,
2007).
3.3. Chronology
Seven AMS radiocarbon dates, determined on plant fragments
and bulk organic matter (Table 1) provide the chronological control
for the Laguna Formosa palynological record. Radiocarbon dates
were calibrated against the Southern Hemisphere curve, SHCal13
(Hogg et al., 2013) using the program Calib Rev. 7.0.2 (Stuiver et al.,
2005). Age-depth models were constructed by Bayesian age-depth
modelling approach using the program “Bacon” (Blaauw and
Christen, 2011).
3.4. Numerical analyses
Percentage pollen data were square root transformed prior to
numerical analyses in order to stabilize their variances. Pollen
zones were determined by CONISS stratigraphical constrained
cluster analysis, and diagrams were drawn using TGView 2.0.4
program (Grimm, 2004). Pollen types used for these analyses were
selected according to their proportions and their ecological
importance, and CYPERACEAE and aquatic taxa were excluded due to
their local over-representation.
To accurately interpret the vegetation changes from LFOR1
sequence, fossil pollen assemblages were compared with 46 surface
pollen samples from the campos region (Mourelle and Prieto, 2012,
2016). Principal Component Analysis (PCA) was performed on
modern and fossil pollen samples, using CANOCO version 4.5 pro-
^
gram (ter Braak and Smilauer, 2003).
In addition, the pollen dispersal capacity of the most frequent
woody taxa present in the pollen surface assemblages from the
Laguna Formosa watershed was analyzed in order to evaluate
which pollen types can be used as reliable riparian forest indicators
and in which proportions they are likely to be represented in the
fossil pollen sequence.
4. Results
4.1. Local modern pollen-vegetation relationship
Woody taxa from riparian forests have mostly zoophilous
pollination and so they produce low quantities of poorly dispersed
pollen grains (Fig. 2). Inside the riparian forest woody pollen rep-
resents ca. 70%, while more than 250 m away from the riparian
forest woody pollen does not reach 2% of the pollen assemblage
(Fig. 2b) (Mourelle and Prieto, 2016). The dispersal capacity of these
pollen grains is determined by ecological characteristics of their
parent plants, such as their position in the forest related to their
water requirements and the height of the shrubs (Fig. 2c), which
has been taken into account to interpret the forest development
reflected by the fossil pollen spectra.
Pollen assemblages from riparian forests (samples 61, 62, 63 and
65) were dominated by woody taxa (55e58%), mainly represented
by MYRTACEAE (up to 25%), hydrophilous taxa (e.g. SALIX CHILENSIS,
CEPHALANTHUS GLABRATUS, SEBASTIANIA/ACANTHOSYRIS), SYAGRUS-TYPE, ALLOPHY-
LUS EDULIS, and MYRSINE (Fig. 2a), the latter being over-represented in
sample 62 (54%). RHAMNACEAE (Scutia buxifolia) and CALLIANDRA (<1%)
are represented only in the riparian forest (Fig. 2c), as well as
DAPHNOPSIS RACEMOSA which is also observed in the wet meadow.
Pteridophytes are mainly represented by POLYPODIUM/MICROGRAMMA
spores (10e20%) (Fig. 2a).
Bunchgrasses (samples 58, 59 and 60) are dominated by POACEAE
(<40%), associated with CYPERACEAE (15e20%), ASTERACEAE SUBF. ASTER-
OIDEAE (15e20%) and ERYNGIUM (<20%) (Fig. 2). Woody taxa are mainly
represented by MYRTACEAE, CELTIS, LITHRAEA/SCHINUS, and hydrophilous
taxa such as CEPHALANTHUS GLABRATUS, PHYLLANTHUS SELLOWIANUS, SALIX CHI-
LENSIS and SEBASTIANIA/ACANTHOSYRIS, reaching up to 20%. CELTIS and
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Table 1
Radiocarbon dates from LFOR1 core.
Sample depth (cm) Uncalibrated age (14C yr BP) Calibrated age weighted average/cal yr BP (2s interval)
05e06 112 ± 37 91 (4e260)
49e51 1220 ± 34 1087 (981e1180)
77e79 2078 ± 39 1997 (1897e2140)
98e99 2936 ± 38 3028 (2886e3162)
112e113 11.250 ± 180 13,061 (12,728e13,382)
115e117 11.922 ± 72 13,686 (13,485e13,951)
156e157 12.453 ± 87 14,517 (14,128e14,982)
a
NSF Arizona AMS Facility.
b n
Conventional date Laboratorio Datacio 14 tedra de Radioquímica, Facultad de Química (UdelaR).
C, Ca
LITHRAEA/SCHINUS are better represented in bunchgrasses than in the
riparian forest (Fig. 2). Pollen from aquatic taxa reaches up to 5%.
Pollen assemblage of the wet meadow vegetation (sample 64) is
similar to bunchgrasses but it contains higher CYPERACEAE pro-
portions (35%) (Fig. 2a).
Grasslands regional pollen dominance is masked in Laguna
Formosa surface sample by woody pollen from riparian forests
(0_LFOR1, Fig. 2a). This forest surrounding Laguna Formosa acts as a
filter limiting the entrance of pollen from grasslands into the
shallow lake.
EUCALYPTUS and PINUS are present in all surface samples. Other
pollen taxa than those not found neither in the floristic surveys nor
in vegetation descriptions for the URRN were recovered in pollen
assemblages, mainly INGA-TYPE, TREMA-TYPE, ALCHORNEA and DODONAEA
VISCOSA (Fig. 2a).
4.2. LFOR1 core
The sediment core consists of dark gray silty clayed from 163 to
113 cm core depth. From 112 to 20 cm depth gray silt with medium-
fine sand, with a distinct sand layer at 112-103 cm and three less
distinct sand interbeds (72-70 cm, 66-64 cm, 55-54 cm), are pre-
sent. The uppermost 20 cm consist of dark gray silt with fine sand
(Fig. 3).
Changes in the organic matter percentages are closely related to
the lithology (Fig. 3). From the bottom core, the organic matter
percentages show increased fluctuating values that ranges from 5%
up to 14% at 124 cm, followed by a decreasing trend up to 3% at
113 cm, that coincides with frequent plant remains in sediments.
From 112 cm to 20 cm depth, organic matter ranged between 0.5
and 2.5%, followed by an increasing trend towards the top core (up
to 8%).
The radiocarbon dates (Table 1), lithology and organic matter
content (Fig. 3) indicate a sedimentary hiatus at ca. 113 cm. As a
consequence the record represents two separate periods: the lower
core section (163-113 cm) encompass from 14,570 to 13,500 cal yr
BP, interval that coincides with the Antarctic Cold Reversal (ACR)
period (Pedro et al., 2015). The overlaying section (112-0 cm) rep-
resents the last 3280 cal yr BP (Table 1; Fig. 3). Before and after the
sedimentary hiatus, radiocarbon dates and sediment stratigraphy
indicate a continuous sedimentation (Table 1; Figs. 3 and 4). Age
ranges were calculated for each period.
4.2.1. Late Pleistocene (from 14,570 to 13,500 cal yr BP)
At 14,570 cal yr BP, pollen assemblages were dominated by
herbs (ca. 55%) associated with trees and shrubs (23%). These as-
semblages were replaced at 14,000 cal yr BP by woody pollen as-
semblages (70%) (Fig. 3). Aquatic herbs reached up to 10% and are
very diverse, well represented by emergent, submerged and
floating plants (Figs. 3 and 5). Algae were only represented by
Zygnemataceae and cyanobacteria mainly by GLOEOTRICHIA-TYPE.
19
Material Laboratory no.
Bulk organic matter AA99174a
Plant remain AA94755a
Plant remain AA99175a
Plant remain AA96779a
Plant remain URU0563b
Plant remain AA94756a
Bulk organic matter AA94757a
Spores were represented by traces of ISOETES (Fig. 5). Total pollen
concentration showed fluctuating values around ca. 100,000 gr/
cm3, and decreased towards the end of this time window (Fig. 3).
Three pollen zones were defined based on CONISS cluster
analysis for this period (Figs. 3, 5 and 6). LFOR1_1 (163-150 cm;
14,570e14,310 cal yr BP) is characterized by the dominance of herbs
(ca. 55%), mainly represented by POACEAE (ca. 45%) associated with
ERYNGIUM (5e10%). ASTERACEAE SUBF. ASTEROIDEAE reached up to 15%. Trees
were manly represented by SALIX CHILENSIS (<20%), shrubs by CELTIS
(<15%), CEPHALANTHUS GLABRATUS (ca. 5%), and SEBASTIANIA/ACANTHOSYRIS
and MIMOSA in traces. RHAMNACEAE and VACHELLIA CAVEN pollen, in traces,
are also present in some samples. Aquatic plants were very diverse,
represented by emergent (mainly CYPERACEAE associated with ECHI-
NODORUS and ALTERNANTHERA), submerged (mainly MYRIOPHYLLUM) and
floating plants (mainly AZOLLA and EICHHORNIA).
LFOR1_2 (150-136 cm; 14,310e14,020 cal yr BP) registered an
increase in trees and shrubs, and co-dominated together with
herbs, mainly due to the maintenance of SALIX CHILENSIS values (<20%)
and to the increase of CEPHALANTHUS GLABRATUS and SEBASTIANIA/ACAN-
THOSYRIS (up to 25%). MYRTACEAE and PHYLLANTHUS SELLOWIANUS became
frequent (<6%) and showed an increasing trend towards the end of
the period, whereas CELTIS decreased. RHAMNACEAE and VACHELLIA CAVEN
pollen are still present. The aquatic plants remained diverse,
although an increase of ECHINODORUS was registered and SAGITTARIA
appears more frequently.
LFOR1_3 (136-113 cm; 14,020e13,500 cal yr BP) is characterized
by the dominance of trees and shrubs, reaching up to 70%, mainly
due to the increase of PHYLLANTHUS SELLOWIANUS that fluctuated around
40% (reaching up to 55%). CEPHALANTHUS GLABRATUS and SALIX CHILENSIS
displayed high values (10e20%), associated with CELTIS (<10%),
SEBASTIANIA/ACANTHOSYRIS (<5%), MYRTACEAE and RHAMNACEAE (<2%).
VACHELLIA CAVEN pollen is no longer present. Aquatic plants exhibited
the same previous characteristics, although POTAMOGETON increased
frequencies and CABOMBA was also observed.
4.2.2. Late Holocene (from 3280 cal yr BP to present)
This period was characterized by a high representation of pollen
of herbs (Figs. 3 and 5). CYPERACEAE was over-represented, reaching
up to 40%. Nevertheless, woody pollen increased gradually towards
the top of the core (from <10% to 25e30%), as well as ASTERACEAE SUBF.
ASTEROIDEAE. Woody pollen, as well as spores from terrestrial plants
(mainly PHAEOCEROS, POLYPODIUM/MICROGRAMMA, and aquatics AZOLLA and
ISOETES), were more abundant and diverse than in the previous time
window. Aquatic herbs showed and inverse trend reaching up to 6%
and were mainly represented by emergent (e.g. TYPHA, ECHINODORUS,
POLYGONUM), submerged MYRIOPHYLLUM) and floating (EICHHORNIA)
plants. Algae (mainly BOTRYOCOCCUS and PEDIASTRUM) and cyanobac-
teria (mainly GLOEOTRICHIA-TYPE) decreased towards the top of the
core. Total pollen concentration values were relatively stable at ca.
10,000 gr/cm3, but increased at the top to 210,000 gr/cm3 (Fig. 3).
Three pollen zones were defined based on CONISS cluster
20 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Fig. 3. Summary pollen and NPPs percentage diagram from LFOR1 core. Lithology, radiocarbon dates, total pollen concentration, organic matter content and pollen zones.
Exaggeration 5.
analysis for this period. Subzone LFOR1_5a/5b was visually estab-
lished to accommodate distinctive changes in the appearance and
the increase of some taxa with ecological significance (Figs. 3, 5 and
7).
LFOR1_4 (112-85 cm; 3280-2270 cal yr BP) was characterized by
the dominance of herbs (60e85%), mainly represented by POACEAE
(40e55%). Minor proportions of ASTERACEAE SUBF. ASTEROIDEAE (<20%)
and ERYNGIUM (<15%) were also well represented. Woody taxa
increased their proportion from 10% to 20% towards the upper
section of the zone, mainly represented by CEPHALANTHUS GLABRATUS
(<4%), PHYLLANTHUS SELLOWIANUS (<3%) and SALIX CHILENSIS (<2%).
LFOR1_5 (85-26 cm; 2270-540 cal yr BP) was also characterized
by the dominance of herbs (ca. 65%). Woody taxa pollen fluctuated
between 25 and 30%, mainly represented by CEPHALANTHUS GLABRATUS
(<20%) and PHYLLANTHUS SELLOWIANUS (<10%). RHAMNACEAE and ALLOPHYLUS
EDULIS (<1%) were also recorded in subzone LFOR1_5b (ca. 940 cal yr
BP). MYRSINE and MIMOSA (<5%), MYRTACEAE and PHYLLANTHUS SELLOWIANUS
(<10%) increased at LFOR1_5b, whereas CEPHALANTHUS GLABRATUS
(<10%) and EPHEDRA TRIANDRA (<1%) decreased.
LFOR1_6 (26-0 cm; 540 cal yr BP-present) was characterized by
pollen assemblages co-dominated by herbs (35e55%) and trees and
shrubs (30e45%), and the increase of ASTERACEAE SUBF. ASTEROIDEAE
(10e25%). ERYTHRINA CRISTA-GALLI and DAPHNOPSIS RACEMOSA were recor-
ded although in low percentages (<1%). CELTIS and SEBASTIANIA/ACAN-
THOSYRIS (<5%) and MYRTACEAE (<20%) increases their relative
abundance, as well as TRIPODANTHUS ACUTIFOLIUS and POLYPODIUM/MICRO-
GRAMMA (<10%), whereas SALIX CHILENSIS decreased (<2%). PINUS and
EUCALYPTUS pollen from alien plants were registered in traces at the
uppermost part.
During the late Holocene, pollen from plants not found in our
own (unpublished data) and other authors' floristic surveys at
URRN (Grela and Brussa, 2003; Brussa and Grela, 2007; Haretche
et al., 2012) were recovered in pollen assemblages, such as INGA-
TYPE, TREMA-TYPE, ALCHORNEA, COMBRETACEAE/MELASTOMATACEAE and DODO-
NAEA VISCOSA (<2%). In addition, pollen from ARAUCARIA was found since
25 cm depth (ca. 510 cal yr BP) (Fig. 7).
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29 21

4.3. Relationship between fossil and modern pollen assemblages

Modern pollen studies carried out at local (Fig. 2) and regional

scale (Mourelle and Prieto, 2012, 2016) were used to compare with
the LFOR1 fossil record, and to study the analogy between modern
and fossil assemblages (Fig. 8). The ordination diagram in Fig. 8
depicts the trajectory of pollen composition changes in relation to
modern pollen assemblages for the two time windows.
The first two PCA-axes accounted for 58.7% of the total variance
(Fig. 8), and split the samples according to their woody pollen
content. As a result, riparian forest surface samples were separated
from grasslands, bunchgrasses and freshwater marshes. On the
other hand, a temporal directionality in the ordination of fossil
samples was noticed both for the late Pleistocene and the late
Holocene periods, where the older samples exhibited higher non-
woody pollen proportions unlike the modern ones.
For the late Pleistocene period, the progressive replacement of
herbs with woody hydrophilous taxa appears to drive the
arrangement of the samples that does not correspond to any
modern analogue. For the late Holocene period, the oldest samples
were dominated by herbs and reflect a partial analogy with
grasslands. The abundance and diversity of woody pollen increased
over time, but their proportions did not reach those observed for
the late Pleistocene. The end of late Holocene period was charac-
terized by pollen assemblages similar to the riparian forest of the
modern samples (Fig. 8).
Despite in each time window woody pollen increases over time,
during late Pleistocene such increase is principally related to the
remarkable high abundance of the hydrophilous taxa, whereas in
the late Holocene is related to the high diversity of pollen types. The
latter determines a different temporal directionality in the ordi-
nation of fossil samples for each time window (Fig. 8).

5. Discussion

5.1. Local modern pollen-vegetation relationship

Riparian forest assemblages were dominated by woody taxa

pollen, represented by hydrophilous, mesophilous and xerophilous
taxa. ASTERACEAE SUBF. ASTEROIDEAE and herbaceous pollen mainly cor-
responds to plants that develop in the understory and to shrubby
taxa located in the edges (Fig. 2).
Since hydrophilous shrubs reach 3e4 m height and grow limited
to water riverbanks, their pollen grains show very low dispersal
capacity and therefore they are mainly found in samples inside the
Laguna Formosa water body (e.g. CEPHALANTHUS GLABRATHUS, PHYLLANTHUS
SELLOWIANUS, MIMOSA and SEBASTIANIA/ACANTHOSYRIS) (Fig. 2c).
C. glabrathus is a profusely branched shrub, and numerous blos-
soms facilitate the arrival of their grains also to bunchgrasses
(Fig. 2c).
Hydrophilous Salix chilensis and mesophilous Sebastiania com-
mersoniana, Acanthosyris spinicens, Blepharocalyx salicifolius (Myr-
taceae), Myrsine laetevirens, Allophylus edulis and Lithraea
molleoides are trees that reach 10e15 m height and constitute the
forest canopy, over which Celtis iguanaea and Tripodanthus acuti-
folius climb. As a result, pollen grains of these taxa not only fall
gravitationally into the forest but they are also dispersed around,
towards the bunchgrasses, the grasslands and the Laguna Formosa Fig. 4. Age-depth model for LFOR1 core for: a) Late Holocene; b) Late Pleistocene.
water body (Fig. 2c). Upper panels depict the MCMC itineration (left); prior (green) and posterior
Scutia buxifolia (Rhamnaceae) and Calliandra tweediei pollen (gray) distributions of accumulation rate (middle); and memory R (right). The bottom
grains drop directly into the ground and they were only found in- panel shows the age-depth model (gray), overlaying the calibrated distributions of the
individual dates (blue). Gray stripped lines indicate the model's 95% probability in-
side the forest (Fig. 2c). Scutia buxifolia is a corpulent tree that tervals. (For interpretation of the references to colour in this figure legend, the reader
reaches up to 10 m height, and C. tweediei is a frequent shrub of the is referred to the web version of this article.)
undersorey that produce large polyades (ca. 100  150 mm).
Daphnopsis racemosa is a small shrub that grows on the forest
 22
}

D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29

Fig. 5. Pollen and NPPs percentage diagram for main palynomorph types from LFOR1 core. Radiocarbon dates, CONISS analysis and pollen zones for each time window. Exaggeration 5.
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Fig. 6. Pollen and NPPs percentage diagram for main pollen types and pollen zones from LFOR1 core for the late Pleistocene time window. Exaggeration 5.
edge close to the littoral of the water body, and so their pollen
grains are only found inside the forest and in the wet meadow. On
the other hand, Celtis ehrenbergiana and Schinus longifolia develop
in the outer forest zone that in front of the bunchgrasses, and so
their grains are easily dispersed towards the bunchgrasses and
grasslands (Fig. 2c).
Bunchgrasses are dominated by herbs (Poaceae and Cyper-
aceae). ERYNGIUM represents E. pandanifolium clumps, and ASTERACEAE
SUBF. ASTEROIDEAE mainly represent Baccharis spp. and Baccharidas-
trum spp. shrubs that develop in the forest edge. Pollen from
aquatic taxa corresponds to freshwater marshes and swampy areas
that develop scattered between bunches. Pollen from woody taxa
represented riparian forest's mesophilous and hydrophilous taxa
(Fig. 2).
Pollen assemblage of the wet meadow vegetation is similar to
those of bunchgrasses but with higher CYPERACEAE proportions,
related to the development of small hydrophilous local commu-
nities in this soils with higher water availability (Fig. 2).
Pteridophyte and bryophyte spores are also present (Fig. 2a).
23
Humid riparian forests understorey promoted the development of
pteridophytes such as Campyloneurum, Pleopeltis and Microgramma
(POLYPODIUM/MICROGRAMMA) and Isoetes (ISOETES). The latter represents
I. weberi, an aquatic emergent/submerged plant that grows on
wetlands or in water riverbanks, as well as in habitats of episodic
inundation, which experiences a dormant phase under dry condi-
tions (Hickey et al., 2009; M. Bonifacino, pers. com. 2009). This
feature explains its relative high frequency also in surface samples
from both grasslands and bunchgrasses (Mourelle and Prieto,
2016).
EUCALYPTUS and PINUS are present in all surface samples and they
represent alien plantations in the region (Fig. 2a).
Pollen grains of TREMA-TYPE, INGA-TYPE, ALCHORNEA and DODONAEA VIS-
COSA could have reached the URRN area by wind-borne dispersal or
river water-borne transported (Fig. 2a). Nowadays, both Trema
micrantha and Inga vera develop along the margin of the Río
Uruguay, more than 300 km westward from URRN, and Dodonaea
viscosa in the Sierra de Ríos low mountains, located 70 km south-
eastward URRN (Fig. 1a and b) (Brussa and Grela, 2007). Moreover,
24 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Fig. 7. Pollen and NPPs percentage diagram for main pollen types and pollen zones from LFOR1 core for the late Holocene time window. Exaggeration 5.
this species constitute the forests and shrublands that develop in
the campos region of southern Brazil (Sobral et al., 2006). In addi-
tion, Alchornea spp. plants are very frequent in Rio Grande do Sul
forests although they are not recorded for the Uruguayan flora (e.g.
Alchornea triplinervia; Sobral et al., 2006). The difficult access to the
URRN region has historically hampered the study of its flora, from
which there is little information in contrast to other Uruguayan
regions. Thus, it cannot be ruled out that, despite these taxa are not
found in our floristic surveys, their parental plants could develop in
close proximity to URRN.
5.2. Paleoenvironmental interpretation
The palynological record of LFOR1 sequence documents the
dynamic balance between grasslands and riparian (gallery) forests
during the late Pleistocene and late Holocene (Fig. 3). The rela-
tionship between modern and fossil pollen assemblages (Figs. 2
and 8) and the pollen dispersal capacity of the most frequent
woody taxa in surface samples (Fig. 2) suggest that despite there
are no-analogs for the interval ca. 14,570 and 13,500 cal yr BP,
hydrophilous woody taxa were widely dominant near Laguna
Formosa since ca. 14,300 cal yr BP. Instead, after ca. 3280 cal yr BP a
partial analogy can be observed, initially between the fossil and the
modern grassland samples, and afterwards with modern riparian
forest samples. During the late Holocene period, grasslands
regionally dominate in URRN, while riparian forests developed near
Laguna Formosa. The gradual increase of the woody taxa pro-
portions in fossil assemblages during the late Holocene, together
with the increased diversity by the incorporation of new taxa,
indicate that the riparian forest changed during this time until
achieving a composition similar to the contemporary time at ca.
540 cal yr BP.
Along the lithological interval dominated by silt-clay (163-
113 cm), dated from 14,600 to 13,500 cal yr BP, shallow and more or
less mesotrophic freshwater habitats develop in URRN, with the
predominance of riparian hydrophilous shrublands and scarce
development of emergent aquatic vegetation in the edges. A sedi-
mentary hiatus at 113 cm depth was probably caused by the Río
Negro southeastward migration (Fig. 1c). Since ca. 3280 cal yr BP,
the meander was isolated from the main channel of Río Negro. The
sand layer at 112-103 cm reflects the early stage of the water body
as an abandoned meander.
The interval with silt with medium-fine sand and three sand
layers interbedded (103-20 cm), dated from 3280 cal yr BP to the
19th Century, reflect a freshwater shallow lake during the late
Holocene suggested by algae, cyanobacteria and the development
of emergent (mainly Cyperaceae), submerged and floating aquatic
vegetation in the edges, together with riparian forest (Fig. 3).
However, lower values of submerged plants than those of late
Pleistocene and coarse-grained lithological levels suggest either
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Fig. 8. PCA distance biplot of the taxa and combined regional modern surface and LFOR1 fossil samples.
high-energy or unstable conditions in the water body which would
have generated more turbid water.
5.2.1. Late Pleistocene (from 14,570 to 13,500 cal yr BP)
Between 14,570 and 14,300 cal yr BP POACEAE dominate pollen
assemblages, reflecting grasslands dominance in URRN. However,
hydrophilous trees and shrubs are also important taxa (Figs. 3 and
6). Although this woody vegetation do not display a high diversity,
their relatively high proportions reflect the development of ripar-
ian hydrophilous shrublands in the littoral of freshwater bodies,
probably similar to those that develop nowadays around some
waterbodies in the campos region. Despite the existence of a robust
regional model of surface pollen-vegetation relationship in the
campos region (Mourelle and Prieto, 2012, 2016), future analysis of
this particular riparian hydrophilous shrublands will probably
allow us to adjust our interpretation about this pollen-vegetation
analogy. However, these pollen assemblages lacking modern ana-
logs should not be dismissed as they have been well documented
for the late glacial period in other regions of the world.
Shrub abundance increased after ca. 14,300 cal yr BP. Sebastiania
commersoniana and S. schottiana represent typical pioneer species
of flooded areas (Budke et al., 2007), that together with Phyllanthus
sellowianus they usually fixe the water riverbanks from an early
stage (Brussa and Grela, 2007). Their increasing trend in the paly-
nological assemblages may be related to the more intense
25
development of hydrophilous shrublands that probably promoted
the fixation of the freshwater riverbanks, thus maintaining shallow,
calm and clear water conditions favorable for GLOEOTRICHIA-TYPE and
Zygnemataceae zygospores development (van Geel and Van der
Hammen, 1978; Carrio  n and Navarro, 2002), and leading to the
increased colonization of floating and submerged aquatic plants,
with a subsequent increase of organic matter content (Fig. 3).
Modern shrublands are not favorable environments for fern and
bryophyte development (Mourelle and Prieto, 2012) which could
explain the absence of these spores in the fossil assemblages.
From 14,000 cal yr BP a further proliferation of hydrophilous
shrublands was recorded, and towards ca. 13,750 cal yr BP shrub-
lands reached their maximum development. The frequent plant
remains suggest gravitational fall either from the leafy plants
around the stagnant or the low flow of water body (Fig. 3).
These lowlands in URRN represent the first direct evidence of
woody riparian vegetation development along the freshwater
bodies in northern campos of RPG during the late Pleistocene. This
vegetation was not only constituted by isolated trees and shrubs as
reported by Behling et al. (2005) for the lowlands in western Rio
Grande do Sul, but also by riparian plant communities. The high
forest diversity suggests that the climate in the northern campos
must have been relatively wet and not so cool between 14,570 and
13,500 cal yr BP to allow such a vegetation development. Despite
this time window coincides with the ACR period (Pedro et al., 2015),
26 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
the palynological analysis appears not to show any evidence of the
ACR cooling event that interrupted the Antarctic deglacial warming
trend in the Southern Hemisphere (Pedro et al., 2015). Laguna
Formosa proxy record, placed between 40
and 20
S, could be
classified into the “unclear” category, where the ACR signals are
weak or even absent (Pedro et al., 2015).
During the last glacial period, pollen records indicate that the
landscape in the northern highlands of Río Grande do Sul were
also dominated by treeless vegetation under a predominantly
cold and dry climate (e.g. Roth and Lorscheitter, 1993; Behling
et al., 2001, 2004; Leonhardt and Lorscheitter, 2010; Jeske-
Pieruschka and Behling, 2012; Scherer and Lorscheitter, 2014;
Spalding and Lorscheitter, 2015). Grasslands dominated the
plateau landscapes and woody taxa were restricted to refuges
with sufficient moisture at lower elevations, suggested as
possible incipient riparian forests along the streams on valleys or
forest close to the margin of canyons, indicated by few pollen
grains of trees or shrubs (Behling et al., 2004; Leonhardt and
Lorscheitter, 2010; Jeske-Pieruschka and Behling, 2012). Such
landscape conditions prevailed through early and mid-Holocene
under warm and dry climates, until starting the initial expan-
sion of Araucaria forests by migration from the gallery forests
along the rivers about 4000 years ago, which indicates a return to
wetter climates (Roth and Lorscheitter, 1993; Behling et al., 2001,
2004; Leonhardt and Lorscheitter, 2010; Scherer and Lorscheitter,
2014). Increased moisture in this phase has been also reported
from other cores of the coastal plains and adjacent areas of Rio
Grande do Sul (Lorscheitter, 2003; Leal and Lorscheitter, 2007). It
should be noted that woody vegetation consists of mostly
zoophilous pollination taxa that produce low quantities of pollen
grains poorly dispersed, and therefore their grains are generally
under-represented in pollen assemblages.
The information provided by both extant and some extinct
vertebrate suggests the dominance of open areas, probably grass-
lands, from 21,000 to 13,000 yr BP in northern Uruguay (Ubilla
et al., 2004). However, according to these authors, the existence
of wooded areas and riparian forests, in permanent lacustrine and
fluvial contexts, should not be dismissed, as indicated by the
occurrence of some extant taxa, such as Lundomys molitor and
Tapirus terrestris, which still exist elsewhere in the continent (Ubilla
et al., 2004). In addition, some mammals recorded in western Rio
Grande do Sul, such as Tapirus and Tayassu pecari, also suggest the
presence of some forested areas during the late Pleistocene (Kerber
et al., 2014).
We found a very striking frequent presence of pollen types that
correspond to some characteristic species of the Chaco and Espinal
floras (Cabrera and Willink, 1973), such as Vachellia caven (VACHELLIA
CAVEN), Celtis ehrenbergiana (CELTIS), Schinus longifolius (LITHRAEA/
SCHINUS) and Scutia buxifolia (RHAMNACEAE). Despite we did not reg-
ister other pollen from taxa that are also characteristic of these
floras (e.g. PROSOPIS), this pollen assemblage could support the hy-
pothesis of Grela (2004) who argues that the xerophytic species
that currently develop near the Río Uruguay may be remnants of
Chaco vegetation that extended to a larger region in the past. Such a
hypothesis is not inconsistent with our interpretation of riparian
hydrophilous shrubland development around freshwater bodies, as
described above. The presence of Prosopis sp. Phyllostylon brasi-
liense and Gleditsia amorphoides wood fragments found in river
deposits in southern Uruguay (site 11, Fig. 1), dated between ca.
13,200 and 12,500 cal yr BP (Ubilla et al., 2017), and of appearance
of some pollen grains of ASTRONIUM and JACARANDA in northern
Uruguay (site 1, Fig. 1) for the same period (Su arez, 2011), also
support this assumption. However, further detailed investigations
are required to obtain more robust results in relation to this
phytogeographic issue.
5.2.2. Late Holocene (from 3280 cal yr BP to present)
Between ca. 3280 and 2270 cal yr BP grasslands and bunch-
grasses regionally dominated in URRN, and riparian forest devel-
oped probably along stream and within river basins, mainly
composed by hydrophilous trees and shrubs, together with other
mesophilous woody taxa (Figs. 3 and 7). During that period, iso-
lated mesophilous trees and shrubs taxa also developed in riparian
forests in southern Rio Grande do Sul, at S~ ao Francisco de Assis
locality (Behling et al., 2005), 260 km northward of Laguna Formosa
(site 2; Fig. 1). The lack of representation of hydrophilous taxa in the
S~
ao Francisco de Assis record could be explained by (1) the low
pollen dispersion capacity of hydrophilous taxa (Fig. 2) and (2) the
distance from the peat bog to the forest developed around Río
Inhacuanda  (600 m), because according to the pollen-vegetation
model (Fig. 2), hydrophilous pollen are hardly represented at dis-
tances >250 m. In addition, Río Inhacuand a is a tributary of the Río
Ibicuí, the main channel located at ca. 7 km from the peat bog. It
may also be possible that the woody plants initially developed
around the main channel (Río Ibicuí) and then migrated progres-
sively to the tributaries (e.g. Río Inhacuanda ). MYRSINE pollen is
observed throughout the whole late Holocene record of LFOR1,
which is in close agreement with Behling et al. (2005) related to the
migration of this genus to these latitudes from the western Rio
Parana  valley during the Holocene.
Between ca. 2270 and 940 cal yr BP, the higher proportions of
CEPHALANTHUS GLABRATUS, PHYLLANTHUS SELLOWIANUS and MIMOSA (e.g.
M. pilulifera and M. cruenta) suggest further development of ripar-
ian forests. These are all riparian species and are very tolerant to
flooding events (Brussa and Grela, 2007). The increased develop-
ment of these hydrophilous shrubs is probably a consequence of
the meander separation from the main river channel. This could
have made easier to these shrubs to colonize and fix the water
riverbanks.
At ca. 940 cal yr BP, riparian forest diversity increased, reflected
by the appearance of RHAMNACEAE, the palm SYAGRUS-TYPE
(S. romanzoffiana) and ALLOPHYLUS EDULIS. Rhamnaceae represents
Scutia buxifolia tree. These species grow inside the riparian forest,
reaching the higher strata (Brussa and Grela, 2007). Scutia buxifolia,
Myrsine laetevirens, Celtis iguanaea and many Myrtaceae are main
riparian forest constituents that increased their abundance,
reflecting the formation of a dense forest. In addition, the lithology
suggests that sand supply to Laguna Formosa ceased at this time,
probably avoided by this dense forest that only allowed the input of
fine sediments during the Río Negro flooding events. In western Rio
Grande do Sul (site 2; Fig. 1), riparian (gallery) forest expanded
forming larger areas along the rivers after ca. 1550 cal yr BP (Behling
et al., 2005).
Since ca. 540 cal yr BP, the riparian forest exhibited a species
composition similar to the current one, when the characteristic
woody taxa reaches the higher values and appeared ERYTRINA CRISTA-
GALLI, CALLIANDRA and DAPHNOPSIS RACEMOSA for the first time in the fossil
spectra (Fig. 7). Taking into account that ERYTRINA CRISTA-GALLI is sub-
represented in modern surface samples (Mourelle and Prieto,
2012), its presence suggest that this tree was highly frequent and
abundant in the riparian forest around Laguna Formosa. The same
fact is interpreted to the only record of CALLIANDRA pollen, which
represents Calliandra tweediei, a frequent undersorey shrub
developed in humid areas and those polyades are really difficult to
find in modern pollen assemblages (Fig. 2c). The presence of these
pollen grains reflects that after ca. 540 cal yr BP the Laguna Formosa
was surrounded by the riparian forest, where its edge was consti-
tuted by hydrophilous plants (Phyllanthus sellowianus, Cepha-
lanthus glabratus and Salix chilensis), followed by mesophilous
species, such as Lithraea molleoides, Schinus longifolia, Allophylus
edulis, Celtis ehrenbergiana, Celtis iguanaea, Myrsine laetevirens,
 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29
Myrsine parvula, Scutia buxifolia and several Myrtaceae. This inter-
pretation is supported by the increased TRIPODANTHUS ACUTIFOLIUS, a
climber species that need a canopy well developed to grow above it.
In addition, POLYPODIUM/MICROGRAMMA spores, represent epiphytes and
plants that develop in the understorey.
From 3280 cal yr BP, climatic conditions should have allowed the
increase of forest abundance and diversity, probably due to higher
relative moisture and water availability. The presence of sand layers
between 1800 and 1200 cal yr BP might indicate frequent and
intense flooding events in URRN. Climate change may have
initialized the expansion of riparian forest into grasslands, followed
by consequently feedback effects such as nutrient accumulation via
increased ground water flow from higher elevations during wet
periods (Silva et al., 2008). In southern campos, in marshes of Paso
Barranca-India Muerta (site 8; Fig. 1), soil water content was vari-
able since ca. 2000 cal yr BP (Mourelle et al., 2015a). In western Rio
Grande do Sul (site 2; Fig. 1), a change in species composition was
recorded between ca. 2270 and 1440 cal yr BP, probably because of
the occurrence wetter conditions, related to higher rainfall and
probably a shorter dry season (Behling et al., 2005). These in-
ferences are also consistent with those interpretations for the
southern Brazilian highland (Behling, 2002). Southward retreat of
the ITCZ in the late Holocene would have increased instability and
amplified the potential for westerly wind anomalies required to
initiate ENSO events (Haug et al., 2001; Koutavas et al., 2006).
Wetter conditions and higher rainfall could have been originated by
increased precipitation over La Plata Drainage Basin related with
the high ENSO amplitude at ca. 2000 cal yr BP (Woodroffe et al.,
2003; Gyllencreutz et al., 2010).
EUCALYPTUS and PINUS represent tree alien species plantations in
the region (Fig. 2a). They were first observed together with
increased organic matter content and total pollen concentration
values (Fig. 3) that could suggest some extent of lake eutrophica-
tion caused by human activities.
5.2.3. Long distance pollen
For the last 3280 cal yr BP, pollen from some taxa that only
develop in the northeastern region of Uruguay were not recorded
(e.g. Xylosma and Citronella), probably due to low pollen produc-
tion. However, pollen from taxa not found in URRN floristic surveys
were recovered in fossil pollen assemblages, mainly INGA-TYPE,
TREMA-TYPE, ALCHORNEA, COMBRETACEAE/MELASTOMATACEAE and DODONAEA
VISCOSA. However, their parental plants are frequent in Rio Grande do
Sul. Even though Laguna Formosa is isolated, it is likely that during
periods of heavy rainfall and associated strong flooding events may
have directly influenced Laguna Formosa and the Río Negro, getting
in contact, as observed in several occasions for other Uruguayan
watercourses (Praderi and Vivo, 1969). However, according to the
proportions and frequencies of these pollen taxa in the late Holo-
cene fossil spectra (Fig. 7) and in the modern samples (Fig. 2a), it
cannot either be ruled out that their parental plants could develop
in close proximity to URRN.
ARAUCARIA was found within the top 25 cm (ca. 510 cal yr BP).
Structural and morphological ARAUCARIA pollen grains characteristics
make it difficult their wind dispersion (Caccavari, 2003). ARAUCARIA
pollen grains found in LFOR1 are not longer than ca. 60 mm in
diameter, which is the maximum pollen length that could be
transported by wind (Faegri and Pijl, 1979) and with relatively poor
preservation. All of this suggests that ARAUCARIA pollen grains could
come from (1) Sierra de Ríos low mountains, where old Araucaria
specimens were found, despite it is unknown whether they are
native or introduced (C. Brussa, pers. com. 2015); (2) the nearest
Araucaria populations, located at 165 and 235 km northeastward
from the study area, in low mountains in the campos of southern
Rio Grande do Sul (>300 m a.s.l.) (Fig. 1), where ARAUCARIA pollen
27
were recovered in a fossil record after 515 cal yr BP, and presumably
correspond to natural populations (Behling et al., 2016); (3) Arau-
caria forests over the planalto sul brasileiro (Fig. 1), in expansion
since ca. 4000 cal yr BP, related to climatic conditions (Behling et al.,
2001, 2004; Jeske-Pieruschka and Behling, 2012) but also to human
seed dispersal (Bitencourt and Krauspenhar, 2006).
6. Conclusions
The palynological record from Laguna Formosa provides evi-
dence that grasslands were regionally dominant throughout late
Pleistocene and the late Holocene periods, but also documents the
dynamic balance between grasslands and riparian forests. The
modern pollen-vegetation relationships and the pollen dispersal
capacity of the most frequent woody taxa improved the vegetation
reconstruction for these two time windows.
The palynological record not only allowed us to identify the
woody flora that developed in northern campos during the late
Pleistocene, mainly composed by hydrophilous trees and shrubs,
but it also represented the first evidence of woody riparian
vegetation development along freshwater bodies during the late
glacial period, being the vegetation constituted not only by iso-
lated individuals. This interpretation is in agreement with in-
formation derived from Uruguayan fossil vertebrate records. The
high forest diversity suggests that the climate must have been
relatively wet and not so cool between 14,570 and 13,500 cal yr
BP to allow such kind of vegetation development. This inter-
pretation appears not to show any evidence of the ACR cooling
event that interrupted the Antarctic deglacial warming trend. In
addition, Laguna Formosa pollen record suggests possible con-
nections between the RPG and the Chaco phytogeographic
province in this latitude during the late Pleistocene as Grela
(2004) had hypothesized.
During the last 3280 cal yr BP, while grasslands were regionally
dominant, riparian forests developed near Laguna Formosa, rep-
resented by hydrophilous and mesophilous trees and shrubs. After
ca. 2270 cal yr BP, the gradual increase in the woody taxa abun-
dance and diversity, reflect that the riparian forest changed during
this time until it achieved a composition similar to the current one
at ca. 540 cal yr BP. Probably higher effective moisture and water
availability that because of increased precipitation over La Plata
Drainage Basin allowed the increase of forest abundance and
diversity.
Pollen from taxa not found in the upper reaches region of the
Río Negro floristic surveys were recovered in fossil pollen assem-
blages, probably as a consequence of overflows from Rio Grande do
Sul. However, it cannot be ruled out that their parental plants could
develop in close proximity to the study area. ARAUCARIA pollen, found
from ca. 510 cal yr BP, would have reached NE Uruguay either by
wind transport or an establishment of small population of Arau-
caria in the area during the late Holocene related to a possible
anthropic contribution in this process.
Acknowledgments
This research was supported by grants of CONICET (PIP 1265)
and Universidad Nacional de Mar del Plata (Exa 15/E550), PEDE-
CIBA and SNI-ANII. We are grateful to Estancia Ana Paula for
permission to work, and D. Panario and R. Bracco for their support
during the field work. We thank R. Bracco for dating sample
URU0563, the (NSF)-Arizona AMS Facility and T. Jull for financial
support for dating. To G.E. Silvestri for the information about
climate of Uruguay, to Brussa and Bonifacino for the botanical in-
formation provided, and to the anonymous reviewers for their
comments and suggestions.
28 D. Mourelle et al. / Quaternary Science Reviews 167 (2017) 14e29

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