Revue de micropaléontologie 68 (2020) 100443

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Revue de micropaléontologie
journal homepage: www.elsevier.com/locate/revmic

Micro/Strat Eastern Medit Special Issue

A multi-proxy approach for reconstructing environmental dynamics since

the mid Holocene in Lake Ismarida (Thrace, N. Greece)
Olga Koukousioura a, * , Katerina Kouli b, Konstantinos Vouvalidis c , Elina Aidona d,
Georgia Karadimou c , Georgios Syrides a
a
Aristotle University of Thessaloniki, School of Geology, Department of Geology, 54124 Thessaloniki, Greece
b
National and Kapodistrian University of Athens, Faculty of Geology and Geoenvironment, Department of Historical Geology-Paleontology, Panepistimioupolis, 15784 Athens,
Greece
c
Aristotle University of Thessaloniki, School of Geology, Department of Physical and Environmental Geography, 54124 Thessaloniki, Greece
d
Aristotle University of Thessaloniki, School of Geology, Department of Geophysics, 54124 Thessaloniki, Greece

A R T I C L E I N F O A B S T R A C T

Keywords: The paleoenvironmental evolution of Lake Ismarida in Thrace (Northern Greece) is revealed by the combined
Benthic foraminifera lithological, micropaleontological (benthic foraminifera, pollen and NPPS), molluscan analyses, magnetic
pollen susceptibility measurement and radiocarbon dating of a 5.8-m long sediment core. The mid Holocene evolution
molluscs
of the lake area is evidenced by the documentation of four sedimentary Units in the core ISMR-2, corresponding to
paleoenvironment
Lake Ismarida
four distinct evolutionary stages: (1) during
5500-3500 cal yr BP the lake area was a shallow marine environment
Thrace characterized by an Ammonia beccarii, small rotaliids, miliolids, Bittium reticulatum and Veneridae spp. assemblage,
Greece marine dinoflagellate cysts, and low magnetic susceptibility values; (2) during
3500-3000 cal BP the environment
is gradually tending to more isolated conditions forming an open lagoon, characterized by marine and euryhaline
fauna and low magnetic susceptibility values; (3) during 3000 cal yr BP, the open lagoon presented a transition to an
oligohaline inner lagoon, characterized by an Ammonia tepida, Haynesina germanica, Aubignyna perlucida, Pirenella
conica, Cerastoderma glaucum and Abra spp. assemblage, sedges and aquatic vegetation. This restricted, entirely
isolated from the sea inner lagoon could be definitely used as the landmark of the Lake Ismaris from Heorodotus,
while describing the march of Xerxes through Thrace in 480 B.C.; (4) since
2000 cal yr BP to the present, the Lake
Ismarida is formed, characterized by fresh-water indicators and aquatic pollen, Pseudoschizaea and high magnetic
susceptibility values. Finally, the progradation of the Filiouris River deltaic deposits resulted to a 4 km wide deltaic
plain between Lake Ismarida and the nowadays coastline. Pollen assemblages record the dominance of a rather rich
deciduous forest in the area with traces of human presence in the lower part of the sequence, whereas the opening of
the plant landscape under the increasing human pressure is evidenced after
3000 cal yr BP. Finally, an open
vegetation pattern, contemporaneous with the retreat of forest vegetation, is evidenced in the area already before
2000 cal yr BP.

1. Introduction
Multidisciplinary studies of sediment cores have been used broadly in
order to reveal the paleoenvironmental evolution, since the last
deglaciation and through Holocene (e.g., Di Rita et al., 2011;
Emmanouilidis et al., 2019; López-Belzunce et al., 2020; Morhange
et al., 2000; Triantaphyllou et al., 2010). All these areas were dynamically
affected by the relative sea level change and the sedimentation pattern,
which were the main factors driven the environmental evolution of the
Holocene coastal plains (Antonioli et al. 2009; Lambeck and Bard, 2000;
Lambeck and Purcell, 2005; Pavlopoulos et al., 2011; Pirazzoli, 2005;
Pirazzoli and Pluet, 1991; Vacchi et al., 2014). This evolution started, as
confirmed, with a rise in sea level causing a marine transgression at the
beginning of Holocene and followed by the progradation of the coastline
during middle Holocene. That resulted to a well recorded environmental
shift to closed environments and the construction of residual lagoons and
lakes, derived by the diminishing rate of the sea level rise, primarily due to
climatic conditions and local tectonics (even anthropogenic activities)
(Amato et al., 2013; Avramidis et al., 2014; Gogou et al., 2007;
Koukousioura et al., 2012, 2019; Lambeck and Chappell, 2001;
Pavlopoulos, 2010; Pavlopoulos et al., 2011; Syrides et al., 2009a;
Triantaphyllou et al., 2009; Vött, 2007; Vouvalidis et al., 2005).

* Corresponding author.
E-mail addresses: okoukous@geo.auth.gr (O. Koukousioura), akouli@geol.uoa.gr (K. Kouli), vouval@geo.auth.gr (K. Vouvalidis), aidona@geo.auth.gr (E. Aidona),
syrides@auth.gr (G. Syrides).

http://doi.org/10.1016/j.revmic.2020.100443

0035-1598/© 2020 Elsevier Masson SAS. All rights reserved.

O. Koukousioura et al.
Several coastal plains related to human activities, settlements, harbors
e.t.c., have been proved as useful archives in order to assess the past
environmental changes (Amato et al., 2020; Belloti et al., 2016; Mariotti
Lippi et al., 2007; Pepe et al., 2013; Russo Ermolli et al., 2014; Sadori
et al., 2010, 2015, 2016). Particularly in the Aegean region where coastal
plains played the most significant role, so that they formed a geographical
cradle for past societies with global cultural impacts, enabled geomor-
phological, paleoenvironmental and sea level reconstructions (Evelpidou
et al., 2010, 2012; Fouache et al., 2008; Ghilardi et al., 2010; Goiran et al.,
2011; Koukousioura et al., 2012; Kouli et al., 2009; Pavlopoulos, 2010;
Pavlopoulos et al., 2007, 2010, 2013; Sidiropoulou et al., 2014; Syrides
et al., 2009a,b; Theodorakopoulou et al., 2009; Triantaphyllou et al.,
2003, 2010, 2016; Vouvalidis et al., 2010a,b). Nevertheless, very few
have been conducted in Northeast Aegean coasts (Ammerman et al.,
2008; Galanidou et al., 2020; Karadimou et al., 2016; Koukousioura et al.,
2012; Pavlopoulos, 2010; Pavlopoulos et al., 2011), although Thrace area
is of great geomorphological and archeological interest. Anthropogenic
activity in the area is confirmed since the Bronze age with the submerged
findings of Glyfada-Mesi Hoard (Simossi, 2009) due to the rise of the sea-
level (Galanidou et al., 2020), and continuous inhabitation since the
Neolithic time (e.g., Ammerman et al., 2008; Efstratiou, 2006; Efstratiou
et al., 1998).
The present study is focused on Lake Ismarida area, located near the
Aegean coasts of Thrace, where anthropogenic activity has been
continuously recorded since the Neolithic period (Arrington et al.,
2016; Bakalakis, 1962; Bakalakis and Sakellariou, 1981), while it has also
been used as a landmark by Herodotus, in Histories, describing the
Fig. 1. (a) Map of Greece pointing to the study area, (b) location map of the broader study area and Lake Ismarida (from Greek Cadastre), (c) geological-geomorphological
map of Filiouris delta plain and Lake Ismarida (from Karadimou et al., 2016), and (d) ISMR-2 core location in the lake area.
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Revue de micropaléontologie 68 (2020) 100443
advance of Xerxes through the Greek territory. Therefore, lithological
determination, micropaleontological, molluscan and pollen analyses,
magnetic susceptibility measurement and radiocarbon dating were all
integrated for the interpretation of a sedimentary sequence at Lake
Ismarida area, in order to reconstruct the paleoenvironmental and
vegetational evolution, and to test the hypothesis of the location and the
diachronic presence of Lake Ismarida at Herodotus’ times.
2. Study area
Lake Ismarida (or Mitrikou) is located in northern Greece, 20 km south
of Komotini city (Thrace), and 3 km inland from the Thracian coasts of the
north Aegean Sea (Thracian Sea), where a chain of lagoons (Elos, Ptelea,
Alyki, Karatza, Xirolimni) is developed from Molyvoti peninsula to the
west (Fig. 1). Lake Ismarida belongs to the Natura 2000 network and is a
Ramsar protected area, while it is the only natural shallow (1 m average
depth) fresh-water lake in Thrace. It is a relatively small elongated lake
displaying a surface of 3.4 km2, and maximum length and width of 2.4 km
and 1.7 km respectively, while it is surrounded by extensive reeds
(Phragmites). Vosvozis River supplies the lake with fresh water,
discharging into its northern part (Fig. 1c). Until 1950 the extra fresh-
water inflow of Vosvozis River, was overflowed to the sea through a
natural channel (Kousouris, 2014). Nowadays the lake is artificially
connected to the sea through a channel, while its water is mainly used for
irrigation.
The lake was formed at the deltaic plain of the Vosvozis and Filiouris
Rivers, bordered on the west by Neogene sediments, forming a low hilly
O. Koukousioura et al.
terrain, and on the east by the alluvial sediments of the Filiouris plain
(Fig. 1c). Quaternary alluvial sediments can be found at the northern part
of the lake (I.G.M.E., 1986; Pisinaras et al., 2007). The basement of the
area consists of the crystalline bedrock of the Rhodope Massif (Mposkos
and Krohe, 2000; Osswald, 1938; Papanikolaou and Panagopoulos,
1981), and eastwards of the lake, Mesozoic metamorphic and
volcanosedimentary series are exposed (I.G.M.E., 1986). The overall
Neogene sediments at the hilly western part of the lake include sands,
silts, sandstones and fossiliferous sandy limestones (Limnocardium,
Congeria) of Paratethys origin, subsequently covered by red-brown
Pleistocene sediments (Syrides, 1998; Syrides et al., 1997).
Uplifting trends are displayed in the northeast Aegean region
(Thrace), and specifically for the adjacent Lafrouda Thrace area a
slight tectonic uplift occurs (-0.3 to -0.5 mm/yr), probably induced by
the trans-tensional tectonic setting of the North Anatolian Fault
(Pavlopoulos et al., 2011). The estimation of sea level changes in the
area can be based on several relevant curves concerning northern
Greece, such as the one of Thermaikos Gulf (Vouvalidis et al., 2005),
Lafrouda Thrace (Pavlopoulos, 2010; Pavlopoulos et al., 2011) and
Samothrace Island (Pavlopoulos et al., 2011; Syrides et al., 2009a).
Nevertheless, the Lafrouda Thrace curve will be taken in account since
in Thermaikos area a subsidence is identified mainly as result of
intense sediment compaction (Pavlopoulos et al., 2011), most
probably due to the loading of the entire Holocene sedimentary cover
(Fouache et al., 2008). On the other hand, in Samothrace the estimated
uplift (
2 mm/yr; Pavlopoulos et al., 2011; Syrides et al., 2009a),
significantly exceeds Lafrouda’s uplift rate. Thus, the curve from
Lafrouda Thrace that corresponds to the study area, suggests a
minimum rate of sea level rise at 0.24 mm/yr during the last 2770
years (Pavlopoulos, 2010).
Lake Ismarida that today is called by the locals as Lake “Mitrikou” or
“Mana” (=mother; relative to its significant value for the life of the area),
was named after the neighboring ancient city of Ismarus, a name-place
reported in Greek mythology. This city was situated near the homony-
mous mountain, that according to Homer, in the Odyssey, was inhabited
by the Cicones and was afterwards destroyed by Odysseus on his way back
to Ithaca. Stravon places the lake between the ancient cities of Ismara and
Maroneia, while Herodotus (7.109) locates it between the ancient cities of
Maroneia and Stryme describing the march of Xerxes through this area in
480 B.C., and subsequently arising a debate about the topology of Thrace
(e.g., Müller, 1975; Tuplin, 2003):
“After crossing the dry channel of the Lissus, Xerxes passed the Grecian
cities of Maroneia, Dicaea, and Abdera, which are in their
neighbourhood, Lake Ismaris between Maroneia and Stryme, and
Lake Bistonis near Dicaea, which receives the waters of two rivers, the
Travus and the Compsatus” (Romm, 2014).
The habitation of the area has been confirmed by the Neolithic
findings of Paradimi (Bakalakis and Sakellariou, 1981), situated at the
north of the lake, and the settlement in Molyvoti peninsula related to
ancient Stryme (Bakalakis, 1967; Arrington et al., 2016). Additionally,
ancient Roman cemeteries have been revealed from the hinderland south
of Lake Ismarida (Terzopoulou, 2000; Loukopoulou et al., 2005;
Arrington et al., 2016).
3. Materials and methods
3.1. Fieldwork
A 5.8-m long sediment core (ISMR-2), was retrieved from the SE shore
of Lake Ismarida (coordinates: 611885,718, 4536536,64) (Fig. 1d), at an
altitude of 0.87 m a.s.l. A Cobra MK1 vibracoring, equipped with 1-m long
and 40 mm in diameter corer was used. Subsequently the core was
transported to the School of Geology of the Aristotle University of
Thessaloniki for further analyses.
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Revue de micropaléontologie 68 (2020) 100443
3.2. Core description and sampling
The systematic description of the ISMR-2 core deposits was
accomplished using as criteria the sediment texture, color (according
to the Munsell soil color chart), grain size and evident depositional
contacts. The sand and mud fractions (% dw) were calculated using wet
sieving with a stainless steel sieve of 63-mm pore size. The systematic
sampling for micropaleontological analysis, was conducted at 2-2.5 cm
interval, or denser whenever was considered necessary, based on the
sediment characteristics. The 51, in total, micropaleontological samples
obtained used both for benthic foraminifera analysis, while 45 of them
where subsampled for palynological analysis, as the sand horizons where
omitted. Furthermore, 191 samples for mollusc analysis at 2.5 cm interval
were obtained from the core.
3.3. Magnetic susceptibility profile
A total of 288 low-field magnetic susceptibility measurements were
carried out in the ISMR-2 core using the MS2 Bartington Susceptibility
meter and a MB2C sensor (ring sensor). The whole core was measured
continuously with a step of 2 cm. The measurements were performed with
the ring sensor on the sealed core tube liners. The whole procedure was
undertaken twice, and average values were calculated.
3.4. Benthic foraminiferal analysis
A total of 51 samples were analyzed for their benthic foraminiferal
content. For each sample 10 g of dried sediment was treated with H2O2 in
order to remove the organic matter and subsequently, washed through a
63 mm stainless steel sieve and dried at 40  C. 300 benthic foraminifera
were obtained whenever was possible, and then determined in species
level, following the generic classification of Loeblich and Tappan (1987,
1994). The total foraminiferal density (FD; number of specimens/g of dry
sediment), and the relative abundances in the benthic foraminiferal
assemblages were calculated, while the diversity indices [species richness
(S) and Shannon-Wiener diversity index (H0 )], were conducted using the
Past.exe 1.23 software package (Hammer et al., 2001).
3.5. Molluscan analysis
Molluscan analysis was realized on 191 selected samples throughout
the whole core. 10 g of dry sediment was treated with H2O2, then washed
into 125 mm sieve and oven dried at 40  C (Karadimou et al., 2016). All
molluscs were selected, identified and counted, while several specimens
were small sized and juveniles, thus they were identified in generic level.
Although shell fragments were present in the majority of the samples,
only complete individuals and characteristic fragments of gastropods and
bivalve valves were considered for further analysis. The total mollusc
density (number of specimens/10 g of dry sediment), and the relative
abundances in the mollusc assemblages were calculated.
3.6. Palynological analysis
In-total 45 out of the 51 micropaleontological samples were
chemically processed for palynological analysis, while 6 samples
originating from sandy horizons where omitted. Subsamples of 5 to 8 g
of dry sediment were spiked with known quantity of Lycopodium spores,
chemically treated with HCl (37%), HF (40%), acetolysed and finally
sieved over a 10 mm sieve, while residues were mounted in silicon oil.
Microscopic analysis was performed at a x400 magnification. During
microscopic analysis all palynomorphs, including microscopic charcoal,
were identified and counted. Pollen identifications were based on Moore
et al.’s key (Moore et al., 1991), Beug’s (2004) key, Reille’s (1992-1998)
atlas and the reference collection of Museum of Paleontology and
Geology, National and Kapodistrian University of Athens (AMPG, NKUA),
O. Koukousioura et al.
while for the NPP’s identification Van Geel et al. (1989, 2003), were used.
Pollen concentration was calculated, while, the samples with concen-
trations below 150 grains/gram (7 samples) were considered barren and
excluded from this study. Percentage pollen diagram was constructed
based on a pollen sum of regional pollen grains, excluding aquatic and
hygrophilous pollen and spores with the use of the program TILIA
(Grimm, 1992). The PDI sum (Centaurea, Cichorieae, Plantago, Polygonum
aviculare type, Pteridium, Ranunculus acris type, Sarcopoterium, Urtica
dioica type) has been used to infer past pastoral activities (Kouli, 2015).
Riverine input has been calculated based on the sum of the erosion
indicating NPPs type 207 and Pseudoschizaea. Finally, charcoal
concentrations of all microscopic fragments bigger than 10 mm were
calculated.
3.7. Radiocarbon dating
Four samples consisting of marine mollusc shells (Cerastoderma
glaucum and Acanthocardia tubercolata) and plant remains (Ruppia
maritima) have been collected from different layers of the core sequence
and used for radiocarbon AMS (14C) dating (Table 1). Samples were
submitted to Beta Analytic (Miami, USA). The conventional radiocarbon
ages have been calibrated after OxCal Ver. 4.3 (Bronk-Ramsey and Lee,
2013), by using MARINE13 curve for marine data (Reimer et al., 2013),
with a local reservoir correction factor (DR) of 149  40 as the average
value for the Aegean Sea.
4. Results
4.1. Core ISMR-2 stratigraphic/lithologic description
The sedimentary record of borehole ISMR-2 comprises mainly of
alternating silts and sands. Four different sedimentological Units were
recognized in the core ISMR-2 deposits (Fig. 2). The lowermost Unit A
extends from 580 cm to 454 cm and consists of dark greenish grey to dark
grey silty sand and sandy mud, with layers of fine to coarse sand,
containing abundant mollusc specimens (average sand and mud content
76.56% and 23.44% respectively). Unit B, from 454 to 334 cm, is
composed by dark greenish grey to dark grey sand and silty sand, with
molluscs and plant remains, containing sand in an average of 81.2 %. Unit
C (334 to 149 cm) is distinguished by greenish grey fine sand with more
silty and clayey layers, containing the mollusc species Cerastoderma
glaucum and plant remains, and presenting an abrupt decrease in sand
content (average 40.5 %). Finally, from 149 cm to the top of the sediment
core, Unit D includes dark brown silty clay, and is characterized by the
absence of faunal remains. This uppermost Unit demonstrates the lowest
sand contents varying between 1.06 and 21.92 %.
4.2. Magnetic susceptibility profile
Fig. 2 illustrates the variation of magnetic susceptibility values along
the core, while all measured values are well correlated with the relative
variations of the sand and mud fractions. From the bottom of the deposit
up to 334 cm (Units A and B), low values, providing an average of
7.15  105 SI for Unit A and 6  105 SI for Unit B respectively, are
observed, with slight fluctuations. Unit C demonstrates progressively
higher magnetic susceptibility values, displaying a maximum of
Table 1
Radiocarbon samples of core ISMR-2 conventional radiocarbon dating and calibrated dates through MARINE13 calibration curve (Reimer et al., 2013), with a local
reservoir correction factor (DR) of 149  40 as the average value for the Aegean Sea.
Lab code Depth (cm) d13C (m) 14C age (yr BP)
Beta-397441 155 1.4 2490  30
Beta-397442 373 14.4 2960  30
Beta-397443 459 13 3150  30
Beta-397444 564 +1.4 5210  30
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Revue de micropaléontologie 68 (2020) 100443
52.2  105 SI at 204 cm core depth. Finally, within Unit D, a significant
increase in magnetic susceptibility is noted and, especially, in the
uppermost part, about the top 80 cm of the core the higher values reach up
to 186.2  105 SI.
4.3. Benthic foraminiferal analysis
Benthic foraminifera assemblages are generally well preserved, when
present. Within Unit D (149 cm to the top depth interval), the 12 samples
studied were derived of foraminifera. A total of 60 benthic foraminiferal
species belonging to 29 genera have been identified in ISMR-2 core
(Fig. 3).
Foraminiferal density (FD) values present a high variability,
fluctuating between 4 and 1752 specimens/g, while the S parameter
and H’ index display intense fluctuations as well (S: 4-30 species/sample,
H’ index: 0.95-2.85). Benthic foraminiferal assemblages are mostly
composed of Ammonia beccarii, A. tepida, Haynesina germanica, elphidiids
(mainly Elphidium complanatum and E. gunteri), other small rotaliids
(mainly Rosalina bradyi) and miliolids (mainly Quinqueloculina seminula
and Q. triangularis). However, the bolivinids which participate in the
microfauna with lower percentages, exhibit not less than 8 species in the
total assemblage of 60 (Fig. 3).
Unit A is characterized by the dominance of Ammonia beccarii
exceeding 50% (ISMR-2 540-542,5) of the total benthic foraminiferal
assemblage, accompanied by mainly large specimens of A. tepida (Fig. 3).
Following, Miliolids comprise in average the 35% of the assemblage,
mainly represented by Q. seminula and Q. stelligera. Small rotaliids
(average 15%) and elphidiids (average 13%), have a significant part in the
microfauna too. The FD parameter (max 213 species/g) and H’ index (max
2.68) present relatively high values.
Within Unit B, A. beccarii values evidence a slight decrease, while A.
tepida percentages are increasing. They both appear with equally large
and small specimens. Miliolids display a significant decrease (average
15%), in comparison with the previous Unit, and small rotaliids and
elphidiids range at the same level. The FD parameter and H’ index still
present relatively high values.
In the subsequent Unit C, a further decrease in A. beccarii is observed,
while A. tepida is reaching up to 57% (ISMR-2 160-162.5) and Ammonia
specimens are mainly small. H. germanica, Aubignyna perlucida and E.
gunteri show both a continuous presence and their higher values (max
34%, 16% and 7.5% respectively) throughout the whole core. In contrast
miliolids, small rotaliids and elphidiids appear with their lower values in
the whole sedimentary sequence. The FD parameter display high values
ranging from 273 to 749 specimens/g, while H’ index ranges at the same
level with the previous Units.
In Unit D no benthic foraminifera fauna has been found.
4.4. Molluscan analysis
Molluscs (gastropods and bivalves) are identified into 148 samples out
of the 191 studied, in ISMR-2 core (Karadimou et al., 2016). Additionally,
in some samples Scaphopoda such as Dentalium sp., fragments of
Echinoidea and Crustacea, as well as plant remains were observed. The
assemblage was relatively rich, including ten gastropod and seventeen
bivalve species. The most abundant species, exhibiting a constant
presence throughout the core, are the Bittium reticulatum, Cerastoderma
2 s (cal yr BP) 2 s (cal yr BC) Marker
2095-1850 145BC-100AD Cerastoderma glaucum
3015-3005 1065-1055 Ruppia maritima
3410-3340 1460-1390 Ruppia maritima
5555-5295 3605-3345 Acanthocardia tubercolata
O. Koukousioura et al.
Fig. 2. Core ISMR-2 lithologic description, radiocarbon dating, vertical variation (dw%) of sand and mud fractions and magnetic susceptibility (MS) profile, within the
identified depositional Units A-D.
Fig. 3. Core ISMR-2 relative abundances (%) of the most common foraminiferal taxa, benthic foraminifera Density (Forams/g), Shannon-Wiener diversity index (H΄),
within the depositional Units A-D.
glaucum, Rissoa spp. and Loripes lacteus, followed by Veneridae spp.,
Mactra sp. and Cerithium vulgatum (mainly in the lower part of the deposit)
(Fig. 4).
In Unit A, the highest number of identified mollusc species (26 taxa)
and individuals is observed reaching 186 specimens/10 g; the macro-
fauna is mainly represented by B. reticulatum exceeding 50% of the
molluscan assemblage in some samples, and Veneridae spp. that present
the higher concentrations of the whole borehole sequence (max 62.8% in
sample ISMR-2 487.5-490), along with Rissoa spp., Caecum trachea,
Cerithium vulgatum, Tellina sp., Tapes sp.; Mytilaster sp., Barbatia sp. and
Pecten sp. participate in the assemblage with minor values (less than 5%).
Dentalium sp., echinoid and crustacean fragments were observed almost
in every sample of this Unit.
Within Unit B, the macrofauna still appears diversified (20 taxa),
although with lower mollusc density (average 36 specimens/10 g),
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consisting mainly of B. reticulatum that ranges at the same levels with the
previous Unit. Similar ranges characterize also the rest of the assemblage,
although with some fluctuations, whereas Cerastoderma glaucum presents
a small concentration (average <3%). Echinoid fragments were observed
in several samples, along with scarce Dentalium sp. and crustacean
fragments.
A significant increase in Pirenella conica (average 39%), is observed
in Unit C, although an abrupt reduction of species (15 taxa) and
individuals (average 21 specimens/10 g) occurs. Cerastoderma glaucum
values also increase (average 11%) presenting small and thin shells,
while Abra spp. appears in the upper part of this Unit. Mactra sp., Rissoa
spp., C. trachea and L. lacteus still participate in the assemblage but with
lower values. Rare echinoid fragments were observed in few samples of
this Unit.
Finally, Unit D lacks any mollusc fauna.
O. Koukousioura et al.
Fig. 4. Core ISMR-2 relative abundances (%) of the most common molluscan taxa, mollusc Density (molluscs/10 g), total Gastropod and Bivalve concentrations (per 10 g),
within the depositional Units A-D.
4.5. Palynological analysis
Out of the 45 samples analyzed, only 38 bared sufficient palynomorph
load to be included in the present study, while 7 subsamples were
characterized as barren. For the samples presented herein pollen
concentration exhibits high fluctuations varying from 47 to 29.500
grains per dry sediment gram, while average concentration is
6.000
grains per gram. Despite the discontinuities, three different local pollen
assemblage zones (LPAZ) were distinguished in the pollen record of core
ISMR-2, based on the abundances of characteristic pollen types (Fig. 5).
LPAZ A (500-465 cm) is characterized by the high arboreal pollen (AP)
abundances (85%). Pinus (45%) and deciduous Quercus (35%) are the
dominant taxa, while Carpinus, Ulmus, Alnus, Fagus and Abies are also
present in lower abundances (Fig. 5). Herb vegetation is mainly
Fig. 5. Core ISMR-2 summary percentage diagram of the palynomorphs within the depositional Units A-D. Fresh water indicators (algae) and marine indicators
(dinoflagellates and foraminifera linings) are presented as concentrations.
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represented by Poaceae (10%). Microscopic charcoal concentrations
reach values of 40,000 particles/gram in the lower part. The marine
dinoflagellate cysts like Spiniferites, Operculodinium, and Lingulodinium,
and foraminifera linings, present high abundances with total marine
indicators reaching more than 58,000 grain/g (Fig. 6).
LPAZ B (352-220 cm) records significantly decreased AP abundances
(45%) as a result of the drop in Pinus and deciduous Quercus abundances.
In contrast, the presence of evergreen Quercus appears slightly increased
compared to LPAZ-A. Chenopodiaceae (25%), Poaceae (15%) and
Cichorieae (15%) are the dominant herb taxa. PDI appears increased,
while microscopic charcoal fluctuates significantly throughout this
interval reaching maxima values of approx. 235,000 particle/g. Marine
dinoflagellate cysts are recorded in lower abundances in this interval,
while foraminifera linings appear more abundant.
O. Koukousioura et al.
Fig. 6. Core ISMR-2 palynological diagram of main pollen types encounted, pollen concentration and microscopic charcoal concentration. PDI (Pollen Disturbance Index)
is after Kouli, 2015.
During LPAZ-C (180 cm-top core) a further decrease in AP (
20%) is
evidenced as a result of the dramatic decrease in all deciduous taxa, while
Pinus is the major tree element especially at the upper part of the core.
Poaceae (25%), Cichorieae (20%) and Chenopodiaceae (20%) are the
main herb taxa, while increased is the abundance of Artemisia (5%).
Coprophilous fungal remains and PDI are increasing. The marine
dinoflagellate cysts are practically absent, while Zygnemataceae appear
in the deposit.
5. Discussion
The combined micropaleontological, palynological and molluscan
analyses with magnetic susceptibility and grain size measurements, and
radiocarbon dating of the core ISMR-2 deposits enabled the recognition of
four different evolutionary stagesof Lake Ismarida during the last5500 years.
From around 5500 and until 3500 cal yr BP (Unit A), shallow marine
conditions prevailed in the Lake Ismarida area that was in direct
communication with the sea. During this time interval a silty sand and
sandy sequence was deposited (Fig. 2), displaying low magnetic
susceptibility measurements and confirming the presence of marine
deposits (Ghilardi et al., 2008; Liu et al., 2012). The microfaunal
composition (Fig. 3) comprises typical species of the infralittoral zone,
such as Ammonia beccarii, A. tepida, other small rotaliids, elphidiids and
miliolids (Brönnimann et al., 1992; Moulfi-El-Houari et al., 1999; Murray,
2006; Sgarrella and Moncharmont Zei, 1993), similar to modern coastal
environments of north Aegean Sea (Dimiza et al., 2016; Koukousioura
et al., 2012). Furthermore, the marine mollusc species, such as Veneridae
spp. and Bittium reticulatum, that dominate the macrofaunal assemblage,
indicate a typical composition of shallow marine environments (Syrides,
2008; Vouvalidis et al., 2010a,b; Evelpidou et al., 2010, 2012; Karadimou
et al., 2016). The same hypothesis is also suggested by the high
abundances of marine dinoflagellate cysts like Spiniferites, Operculodinium
and Lingulodinium machaerophorum along with foraminifera test linings
(Fig. 5; LPAZ A) (Elshanawany et al., 2010; Kouli et al., 2009; Morzadec-
Kerfourn, 1992). The occurrence of the halophytic Chenopodiaceae
represent near shore vegetation, while Cyperaceae, pollen from aquatic
plants such as Sparganium and the fresh-water indicators Pseudoschizaeae
and type 353 (Van Geel et al., 1989), should be considered transported
into the marine environment by local streams or rivers. The micro- and
macrofauna composition and the palynological data in combination with
the minimum magnetic susceptibility values confirm the occurrence of a
shallow marine environment, in the present-day Lake Ismarida area. The
lake area was flooded by the sea as a result of the maximum marine
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Revue de micropaléontologie 68 (2020) 100443
transgression, according to the proposed for Thrace, sea level curve for
Thrace (Pavlopoulos, 2010; Pavlopoulos et al., 2011), constituting a
shallow marine environment from 5500 to 3500 cal yr BP.
During the same interval, a rather rich and diverse deciduous forest is
recorded, where even though deciduous Quercus was dominant, Carpinus
was abundant and Ulmus, Fraxinus, Corylus, Tilia, Cornus, and Rosaceae
also flourished. This is in accordance with regional pollen records that
register the development of mixed oak forests at low and mid-altitudes in
the area of southern Balkan (Lazarova et al., 2011; Marinova et al., 2012;
Marinova and Ntinou 2018; Masi et al., 2018; Sadori et al., 2015).
Mountainous forests were also well developed and both altitudinal zones
of conifers (Pinus, Abies, and some rare Picea) and Fagus forests are
represented in the pollen record. Herb vegetation is rather restricted and
mainly consists of Poaceae and Chenopodiaceae. Human presence in the
area is featured by the occurrence of several apophytes (ruderal taxa) and
coprophilous fungi, while no evidence of cereal cultivation in the vicinity
of the coring site has been detected. The low values of PDI (pollen
disturbance index; Kouli, 2015) and the lack of cereal cultivation indicate
low human pressure in the surroundings of Lake Ismarida, most probably
due to the location of the lake far from settlement areas (Efstratiou et al.,
1998; Kouli, 2015; Mercuri et al., 2019).
In the next evolutionary phase of Lake Ismarida, lasting from 3500 to
3000 cal yr BP, the area is gradually becoming more isolated from the sea,
as indicated by the gradual appearance of mesohaline faunal indicators
(Unit B). The benthic foraminiferal fauna is still mainly represented by
marine species, although the percentage of large and small Ammonia spp.
specimens, suggest mesohaline conditions (Evelpidou et al., 2010, 2012;
Koukousioura et al., 2012). Additionally, the association of the marine
with the euryhaline species H. germanica, A. perlucida and E. gunteri,
indicative of mesohaline conditions, compose a typical open lagoonal
assemblage affected by the sea (Frontalini et al., 2013; Koukousioura
et al., 2019; Triantaphyllou et al., 2010). Similar conditions are confirmed
by the macrofauna that is dominated by the strongly fluctuating
abundances of the marine euryhaline species Bittium reticulatum and
Mactra sp. At the same time, the marine species of Veneridae, that were
dominating during the previous stage, are becoming less important. The
overall faunal composition in combination with the low magnetic
susceptibility values, suggests an open lagoon environment in good
communication to the sea with occasional fresh-water inputs. During the
same time interval, the operation of the open lagoon system in Lake
Ismarida coincides with the existence of the already established open
paleo lagoon of Lafrouda Thrace (Koukousioura et al., 2012), westwards
to the Thracian coast.
O. Koukousioura et al.
At around 3000 cal yr BP, the faunal, floral and magnetic susceptibili-
ty data suggest the beginning of the prior evolutionary stage lagoon’s,
entire isolation from the sea. This significant shift in Lake Ismarida’s
conditions, from open to restricted environmental conditions, can be the
cumulative result of the reduction of the sea level rise rate and the
constant deltaic prolongation of Filiouris River, that could generate the
construction of a beach barrier in the study area. During the same interval,
the adjacent Lafrouda Thrace lagoon shows similar evolution, displaying
a gradual change from an open lagoon system to a closed one, a shift also
identified in other Aegean coastal plains during different time spans (e.g.,
Goiran et al., 2011; Triantaphyllou et al, 2003, 2016; Vouvalidis et al.,
2010a,b). This paleoenvironment succession recorded in the study area
during this interval is in good agreement with the suggested sea level
curve for Northeast Aegean (Pavlopoulos, 2010; Pavlopoulos et al.,
2011).
The faunal (benthic foraminifera and mollusc) composition of Unit C
(3000 to 2000 cal yr BP; Fig. 4,5), provides evidence for the establishment
of a restricted oligohaline lagoonal environment, since foraminifera
assemblages are characterized by euryhaline and low salinity tolerant
species. A. tepida is an opportunistic taxon tolerant to a wide range of
salinity and temperature lagoonal and near-shore environments (Almogi-
Labin et al., 1992; Coccioni, 2000; Frontalini et al., 2009; Jorissen, 1988;
Koukousioura et al., 2012; Melis and Violanti, 2006), that produces
smaller and thin-walled tests in low salinity and pH conditions, as
observed from worldwide brackish and estuarine assemblages (e.g., Alve
and Murray, 1994; Debenay et al., 1996; Frontalini et al., 2009;
Koukousioura et al., 2012; Melis and Violanti, 2006; Murray, 1991; Van
Der Zwaan, 2000). Also, Haynesina germanica, is an euryhaline significant
shallow-water inhabitant (eg., Jorissen, 1988; Murray, 2006), which
favors shallow silty lagoonal environments (e.g., Hottinger et al., 2001).
In this time interval, the association of the small-test sized Ammonia
specimens observed in the assemblage, with H. germanica and A. perlucida
points to restricted salinity conditions (Koukousioura et al., 2012).
Molluscan fauna is characterized by smaller assemblages and the intense
presence of euryhaline species such as Pirenella conica, Cerastoderma
glaucum and Abra spp. P. conica tolerates oligohaline shallow lagoonal
conditions, and develops dense populations preferring the edge of water
(Paperna and Overstreet, 1981), while Abra spp. favors silty lagoonal
oligohaline conditions (Fabbrocini et al., 2008; Gontikaki et al., 2003).
Furthermore, C. glaucum lives in large numbers in shallow lagoons, while
salinity and temperature influence shell size and thickness, resulting in
smaller and thinner shells in oligohaline conditions (Eisma, 1965;
Tarnowska et al., 2009). The molluscan association of this interval in
combination with the small and thin shells of C. glaucum and the decrease
in abundance of the marine euryhaline species (Bittium reticulatum and
Veneridae spp.), indicate inner lagoonal conditions with fluctuating
salinity (Kevrekidis et al., 1996; Koutsoubas et al., 2000; Nicolaidou et al.,
1988; Nossier, 1986; Malham et al., 2012), similar to the present-day
inner coastal lagoons of the area (Elos, Ptelea, Alyki; Syrides, unpublished
data). The overall aquatic palynomorph assemblage (Fig. 5) reflects a
complex depositional environment with both marine and fresh-water
inputs. The abundance and the concentration of dinoflagellate cysts and
foraminifera linings are decreased, in comparison with previous
evolutionary stages, while Cyperaceae, type 353 and type 207 are also
encountered. The palynomorph record further supports the establish-
ment of isolated environmental conditions (Kouli et al., 2009;
Triantaphyllou et al., 2010), as suggested by foraminiferal and mollusc
analysis and magnetic susceptibility measurements. The establishment of
this oligohaline inner lagoon is confirmed after 3000 cal yr BP, although it
was still slightly affected by the sea possibly due to wave overflows over a
closed beach barrier. The restricted water body, entirely isolated by the
sea, could have easily been characterized as a lake, proving correct its use
as a landmark by Heorodotus in 480 B.C., while describing the march of
Xerxes through Thrace. During this interval an opening of the plant
landscape is evidenced by the decrease of AP as a result of the opening of
the forest vegetation and a small retreat in mountainous forests. The
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Revue de micropaléontologie 68 (2020) 100443
mixed deciduous oak forests are neither as dense nor as diverse as before,
while Mediterranean woodland with evergreen Quercus, Pinus, and
Juniperus are becoming a more significant part of the landscape. The
retreat of deciduous forest elements coincides with increased micro-
charcoal concentrations indicating the occurrence of significant regional
fire episodes during the same interval. A smaller retreat is evidenced in
upland forest vegetation. The retreat of deciduous forest and the
expansion of sclerophyllous vegetation is in agreement with patterns
observed in other pollen records (Athanasiadis et al., 2000; Masi et al.,
2018; Panagiotopoulos et al., 2013) as a result of the continuous human
pressure to the ecosystems. The increased abundances and high diversity
of ruderal herb taxa (e.g. Plantago, Polygonum aviculare, Sanguisorba,
Rumex; Bottema and Woldring, 1990; Kouli et al., 2018), and the
amplified PDI values are indicative of increased human presence and the
implementation of grazing activities in the area. This is further supported
by the increased abundances of Cichorieae (Florenzano et al., 2015) and
the presence of coprophilous fungal and parasite remains (Florenzano and
Mercuri, 2012; Kouli, 2012; Van Geel et al., 2003). In addition, repeated
erosional episodes, probably connected with the retreat of forest
vegetation and related human activities in the basin, are detected by
the erosion indicator fungal spore type 207 (Fig. 6). Finally, the amplified
Chenopodiaceae abundances represent the expansion of halophytic
vegetation around Ismarida lagoon during paleoenvironmental Unit B.
During the last evolutionary stage (Unit D), the study area is entirely
isolated from the sea and a fresh-water lake is formed. Lake Ismarida
seems to take its final form at about 2000 cal yr BP. The sedimentary
sequence consists of silty clay deposits (the highest mud values of the
whole sequence). It is characterized by high magnetic susceptibility
values, and a total absence of faunal remains (benthic foraminifera and
molluscs) due to the unfavorable salinity conditions. A rather unstable
depositional environment is indicated by the pollen spectra that are rich
in non-pollen palynomorphs (NPP’s) (Fig. 5). Fresh-water indicators and
aquatic pollen are encountered, while marine dinoflagellate cysts are
absent and foraminifera linings scarce. At the same time increased stream
discharges are reflected, by to high abundances of Pseudoschizaea. The
lack of fauna (foraminifera/molluscs) and the outcome of the palynolog-
ical analysis point to a mainly fresh-water depositional environment with
possible river discharges, even though ephemeral connection with the
marine environment existed. Finally, the progradation of the Filiouris
river deltaic deposits resulted to a 4 km wide deltaic plain between Lake
Ismarida and the nowadays coastline.
The retreat of forest vegetation becomes more evident after 2000 BP,
during LAPZ indicating the establishment of an open landscape in the
surrounding area. Pinus represents the major part of tree vegetation, while
Quercus and other deciduous taxa are scarce. The opening of the
vegetation landscape is evidenced in most regional pollen records and it is
associated with a considerable expansion of human indicator taxa
(Athanasiadis et al., 2000; Kotthoff et al., 2008; Masi et al., 2018;
Panagiotopoulos et al., 2013) Unfortunately, pollen concentration is
extremely low and the plant landscape during this period is not clear.
6. Conclusions
Lake Ismarida, the only existing natural shallow fresh-water lake in
Thrace, represents a dynamic environment that has undergone several
different evolutionary stages during the last 5500 years, as clearly
indicated by the multi proxy study of the ISMR-2 core deposits.
During
5500 to
3500 cal BP, Lake Ismarida was in direct
communication with the sea and shallow marine conditions prevailed,
characterized by an Ammonia beccarii, small rotaliids, miliolids,
Veneridae spp. and Bittium reticulatum benthic assemblage, along with
marine dinoflagellate cysts, and low magnetic susceptibility values.
In-between
3500 and 3000 cal BP, Lake Ismarida area is gradually
tending to more isolated conditions with distinct open lagoonal features,
still in communication with the sea and occasional fresh-water inputs, as
indicated by the appearance of the mesohaline faunal indicators and the
O. Koukousioura et al.
low magnetic susceptibility values. The benthic assemblage is still
represented by marine species, but in association with the euryhaline
species Haynesina germanica, Aubignyna perlucida, Bittium reticulatum and
Mactra sp.
At around 3000 cal yr BP, the open lagoon of Lake Ismarida
demonstrates a transition to an oligohaline inner lagoon, entirely
isolated from the sea, due to the reduction of the sea level rise rate and the
constant deltaic prolongation of Filiouris River. The major characteristics
of the inner lagoon include an Ammonia tepida, H. germanica, A. perlucida,
Pirenella conica, Cerastoderma glaucum and Abra spp. benthic assemblage,
with sedges and aquatic vegetation. This restricted, entirely isolated from
the sea inner lagoon, has been used as a landmark of the Lake Ismaris from
Heorodotus, while describing the march of Xerxes through Thrace in 480
B.C.
Since 2000 cal yr BP to the present day, the fresh-water Lake Ismarida
formed, and supplied with fresh-water discharges, as indicated by the
high magnetic susceptibility values, the fresh-water indicators and
aquatic pollen in combination with the Pseudoschizaea. Finally, the
progradation of the Filiouris River deltaic deposits resulted to a 4 km wide
deltaic plain between Lake Ismarida and the nowadays coastline.
Pollen assemblages record the dominance of a rather rich deciduous
forest in the area with traces of human presence in the lower part of the
sequence, whereas the opening of the plant landscape under the
increasing human pressure is evidenced after
3000 cal yr BP. Finally,
an open vegetation pattern, contemporaneous with the retreat of forest
vegetation, is evidenced in the area already before 2000 cal yr BP.
The environmental evolution of Lake Ismarida during the last 5500
years provides significant data for the coastal processes, sea level changes,
vegetation and the human presence and contributes to the interpretation
of past historical events, in this little-known area of Northern Greece, the
Thracian coastal plains.
Declaration of interests
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
The authors would like to thank the “Molyvoti Project” headed by
Prof. Nathan Arrington, Department of Art and Archeology, Princeton
University, USA, for the financial support of the fieldwork and AMS
datings. Furthermore, the constructive criticism of two anonymous
reviewers is greatly appreciated.
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