diatomaceous sedimentation in late neogene lacustrine basins of western macedonia, greece

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ORIGINAL PAPER Diatomaceous sedimentation in late Neogene lacustrine basins of western Macedonia, Greece R. B. Owen R. W. Renaut M. G Stamatakis Received: 31 August 2009 / Accepted: 19 January 2010 / Published online: 4 February 2010 Ó Springer Science+Business Media B.V. 2010 Abstract Several lacustrine basins were formed by late Miocene tectonic processes in western Macedo- nia, Greece. These were occupied by a series of lakes and wetland swamps during the late Miocene and early Pliocene giving rise to diatomaceous clay- stones, siltstones and diatom-bearing tufa deposits. Diatoms are rare or fragmentary in sandstones and chemical sediments, and are absent in conglomerates. The Fe 2? -rich phosphate minerals anapaite and vivianite are present in parts of the diatomaceous siltstones. Six major diatom groups were identified using cluster and correspondence analyses, each of which tends to be associated with a particular lithofacies. The floras are indicative of mildly acidic to alkaline water of varying depths. The dominant taxa include Aulacoseira ambigua, A. distans, Cyclotella iris and several of its varieties, C. aegeae, C. castracanei, C. elymaea, C. ocellata, and two unidentified Cyclotella. Fragilariaceae are common, consisting mainly of Fragilaria bituminosa, Pseu- dostaurosira brevistriata, P. zeilleri, Staurosira construens, Staurosirella leptostauron, and S. pin- nata. Locally, chysophycean stomatocysts are com- mon, suggesting more oligotrophic conditions. Nine diatom stages are recognized in the Lower Neogene Series at Kariditsa (Kozani basin), reflecting shallow swamps, small alkaline lakes, and terrestrial settings. The Upper Neogene Series rocks are characterized by Mg-rich carbonates such as hydromagnesite, huntite and magnesite. A detailed stratigraphy was developed for Mio-Pliocene deposits in the Florina basin at Klidi. Parts of this sequence show clear cyclicity in both the sediments and the diatom floras, with shallow-water diatoms repeatedly giving way to taxa indicative of deeper conditions. Larger-scale, shal- low-deep cycles are also present and may be related to precession- and eccentricity-forced climate change. Keywords Climatic change Á Diatoms Á Lacustrine basins Á Greece Introduction This study involves investigation of late Miocene and Pliocene diatomaceous deposits in western Macedo- nia, Greece. Several researchers have previously examined Neogene diatom floras in southeastern R. B. Owen (&) Department of Geography, Hong Kong Baptist University, Kowloon Tong, Hong Kong e-mail: [email protected] R. W. Renaut Department of Geological Sciences, University of Saskatchewan, Saskatoon, SK S7N 5E2, Canada e-mail: [email protected] M. G Stamatakis Department of Geology, University of Athens, Panepistimiopolis, Ano Ilissia, 157 84 Athens, Greece e-mail: [email protected] 123 J Paleolimnol (2010) 44:343–359 DOI 10.1007/s10933-010-9409-5

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ORIGINAL PAPER

Diatomaceous sedimentation in late Neogene lacustrinebasins of western Macedonia, Greece

R. B. Owen • R. W. Renaut • M. G Stamatakis

Received: 31 August 2009 / Accepted: 19 January 2010 / Published online: 4 February 2010

� Springer Science+Business Media B.V. 2010

Abstract Several lacustrine basins were formed by

late Miocene tectonic processes in western Macedo-

nia, Greece. These were occupied by a series of lakes

and wetland swamps during the late Miocene and

early Pliocene giving rise to diatomaceous clay-

stones, siltstones and diatom-bearing tufa deposits.

Diatoms are rare or fragmentary in sandstones and

chemical sediments, and are absent in conglomerates.

The Fe2?-rich phosphate minerals anapaite and

vivianite are present in parts of the diatomaceous

siltstones. Six major diatom groups were identified

using cluster and correspondence analyses, each of

which tends to be associated with a particular

lithofacies. The floras are indicative of mildly acidic

to alkaline water of varying depths. The dominant

taxa include Aulacoseira ambigua, A. distans,

Cyclotella iris and several of its varieties, C. aegeae,

C. castracanei, C. elymaea, C. ocellata, and two

unidentified Cyclotella. Fragilariaceae are common,

consisting mainly of Fragilaria bituminosa, Pseu-

dostaurosira brevistriata, P. zeilleri, Staurosira

construens, Staurosirella leptostauron, and S. pin-

nata. Locally, chysophycean stomatocysts are com-

mon, suggesting more oligotrophic conditions. Nine

diatom stages are recognized in the Lower Neogene

Series at Kariditsa (Kozani basin), reflecting shallow

swamps, small alkaline lakes, and terrestrial settings.

The Upper Neogene Series rocks are characterized by

Mg-rich carbonates such as hydromagnesite, huntite

and magnesite. A detailed stratigraphy was developed

for Mio-Pliocene deposits in the Florina basin at

Klidi. Parts of this sequence show clear cyclicity in

both the sediments and the diatom floras, with

shallow-water diatoms repeatedly giving way to taxa

indicative of deeper conditions. Larger-scale, shal-

low-deep cycles are also present and may be related

to precession- and eccentricity-forced climate

change.

Keywords Climatic change � Diatoms �Lacustrine basins � Greece

Introduction

This study involves investigation of late Miocene and

Pliocene diatomaceous deposits in western Macedo-

nia, Greece. Several researchers have previously

examined Neogene diatom floras in southeastern

R. B. Owen (&)

Department of Geography, Hong Kong Baptist

University, Kowloon Tong, Hong Kong

e-mail: [email protected]

R. W. Renaut

Department of Geological Sciences, University of

Saskatchewan, Saskatoon, SK S7N 5E2, Canada

e-mail: [email protected]

M. G Stamatakis

Department of Geology, University of Athens,

Panepistimiopolis, Ano Ilissia, 157 84 Athens, Greece

e-mail: [email protected]

123

J Paleolimnol (2010) 44:343–359

DOI 10.1007/s10933-010-9409-5

Europe and also in neighboring Turkey. Ognjanova-

Rumenova (2000) outlined the general biostratigra-

phy of diatoms in the Balkans, noting that the oldest

lacustrine floras date to the middle Miocene. There

also have been many site-specific studies, particularly

in Bulgaria (Temniskova-Topalova and Ognjanova-

Rumenova 1988; Ognjanova-Rumenova and Popova

1996; Temniskova et al. 1996; Temniskova-Topalova

et al. 1996; Ognjanova-Rumenova 2001, 2004;

Ognjanova-Rumenova et al. 2008). Ognjanova-

Rumenova (2005) presented the Neogene diatom

stratigraphy of a core from the Bitola basin in the

Former Yugoslav Republic of Macedonia (FYROM),

which lies to the north of the present study area.

Rehakova (1980) and Ognjanova-Rumenova and

Vass (1998) have documented the diatoms in Neo-

gene lacustrine deposits in Slovakia, with the latter

authors reporting dominance by several varieties of

Cyclotella iris Brun & Herib. and many Fragilaria

species. Servant-Vildary et al. (1986) and Gurel and

Yildiz (2007) have carried out similar investigations

of Miocene and Pliocene diatoms in Turkey.

Several investigations have focused on taxonomic

aspects of the fossil floras in the region. Examples of

such studies include the work of Ognjanova-

Rumenova et al. (1990) on Fragilaria and Ognjano-

va-Rumenova (2005) on C. iris and its varieties.

Ecomomou-Amilli (1987, 1999) and John and

Economou-Amilli (1990) have presented taxonomic

descriptions of Greek fossil diatoms. Diatom inves-

tigations of single samples from the study area

reported in this work (Ptolemeis and Kozani basins,

Fig. 1) include a taxonomic listing by Gersonde and

Velitzelos (1977) and a description of a new species,

Cyclotella elymaea Ec.-Amilli (1991.

This investigation is based on diatomaceous sed-

iments collected from quarries and natural outcrops

in the Greek Macedonian Florina-Ptolemais-Kozani

basin (Fig. 1a). This constitutes part of the 180-km-

long Pelagonia graben, which extends into the

FYROM and was formed after the Savanian orogeny

and early Miocene peneplanization of the Balkans

(Dumurdzanov et al. 1997). Various parts of the basin

contained lakes from the middle Miocene through to

the end of the Pliocene and perhaps into the

Pleistocene (Dumurdzanov et al. 2004). This study

focuses on the upper Miocene and Pliocene sediments

of the Florina and Kozani basins, with the major aims

of (1) determining the diatom floras; (2) establishing

a high-resolution diatom stratigraphy; and (3)

a bFig. 1 Location and

geology of the Florina-

Ptolemais-Kozani Basin.

a Geographic extent,

location and generalized

stratigraphic section after

Ehlers (1960) in Steenbrink

et al. (2006). b Detail of the

Kozani Basin showing

geology and sampling

locations

344 J Paleolimnol (2010) 44:343–359

123

combining lithofacies and diatom data in order to

reconstruct the paleolimnological conditions that

prevailed in the area during the late Miocene and

Pliocene.

Geological background

The Miocene basins of Greece are usually bounded

by major normal faults while minor faults affect basin

infills (Ioakim et al. 2005; Diamantopoulos 2006).

The location of the NNW–SSE chain of lake basins in

the study area (Florina, Ptolemais, Kozani) is shown

in Fig. 1a. Neogene sediments are widespread within

this tectonic graben, which developed in response to

late Miocene NE–SW extension in the westernmost

zone of the Internal Hellenides (Brunn 1956). Late

Pleistocene NW–SE extension fractured the area into

the Florina, Ptolemais and Servia (Kozani) Basins

(Pavlides and Mountrakis 1987), which extend over a

distance of about 120 km and lie between 300 and

700 m above sea level, adjacent to mountains that

rise to about 2,000 m. The basement includes meta-

morphic rocks, limestones, dolostones, flysch and

ophiolites (Dermitzakis and Papanicolaou 1981).

Steenbrink et al. (2006) have described several

quarry outcrops in the region, noting that about

600 m of lake sediments were laid down during the

Upper Miocene through Lower Pliocene. These

consisted of intercalated lignites, alluvial sands and

conglomerates that occur in four lithostratigraphic

units (Fig. 1a). Shallow lacustrine diatomaceous

deposits and lignites formed the dominant compo-

nents of the Komnina and Ptolomais Formations and

were related to precession-induced climate cycles.

The general environmental setting of the lignites have

been noted by Christanis (2004) with Oikonomopo-

ulos et al. (2008) providing detailed characteristics of

the lignites at Achlada (Florina Basin), close to Klidi

(Fig. 1a). Stamatakis (2004) and Koukouzas et al.

(2009) have documented the detailed mineralogy and

geochemistry of parts of the lacustrine clayey diat-

omites at Vegora and Komnina (Ptolemais Basin),

with Koukouzas (2007) providing descriptions of the

mineralogy of lignites at Achlada (Florina Basin).

Two distinct stratigraphic units have been recog-

nized in the Kozani Basin: a) the Lower Neogene

Series (LNS), and b) the Upper Neogene Series (UNS)

(Fig. 1b) (Anastopoulos and Koucouzas 1969). The

LNS consists of up to 400 m of marlstones with rare

lignite and chert intercalations (Stamatakis et al.

1989). The UNS includes Mg-rich carbonates, marl-

stones, marly diatomites, claystones and poorly com-

pacted ash-tuffs (Stamatakis et al. 1989; Stamatakis

1994, 1995; Calvo et al. 1995; Hall and Stamatakis

2000). Based on botanical evidence, Velitzelos and

Knobloch (1986) reported the age of the UNS in the

southern part of the basin as Late Miocene.

Methods

Samples were collected from seven outcrops of

varying thickness (1–37.3 m) in the Kozani and

Florina Basins (Fig. 1). Two of the longer sections at

Kariditsa (Kozani) and Klidi (Florina) were examined

in greater detail with samples for diatom analyses

collected at 10–15-cm intervals and at lithological

boundaries. Carbonates were removed from the

sediments by adding 10%HCl to about 1 cm3 of

sample, followed by washing in distilled water. This

process assisted with disaggregation of the material.

Aliquots of suspended diatoms and sediment were

then dried on cover slips, prior to mounting on smear

slides with Naphrax. The diatoms were examined

using an Olympus microscope with phase contrast

and brightfield illumination at 1,0009 magnification.

Several diatom identifications, particularly for Cy-

clotella, were confirmed using a LEO 1530 Field

Emission Scanning Electron Microscope.

The various species were mainly identified using

the works of Heribaud (1893), Ehrlich (1966), Gasse

(1980), Serieyssol (1980, 1984), Krammer and Lange

Bertalot (1986, 1988, 1991a, b), Loginova et al.

(1990), Ognjanova-Rumenova et al. (1990), Round

et al. (1990) and Ognjanova-Rumenova (1996). A

minimum of 400 diatoms was counted along transects

on each slide, except where diatoms were rare and all

diatoms were tallied. Diatoms were included in the

counts if there was at least half of the specimen

intact, which may have resulted in some underesti-

mation of the more fragile species. Apices were

counted and divided by two in the case of long thin

taxa such as Synedra in order to minimize errors due

to their fragility. Visual estimates were made of the

percentage of the carbonate-free sample that con-

sisted of diatoms (total intact ? fragmentary).

Diatom assemblages and sampling sites were dis-

tinguished using XLSTAT (Addinsoft) to carry out

J Paleolimnol (2010) 44:343–359 345

123

Agglomerative Hierarchical Cluster Analyses (AHC).

Pearson Dissimilarity and Unweighted Pair-group

Mean Averaging (UPGMA), using all diatom taxa,

were adopted after exploring several AHC techniques.

Dendrograms were generated for both samples and

species (Fig. 5b, d). Only taxa that comprised C5% of

the flora in at least two samples, or C10% in at least one

sample (43 taxa), are shown in the species dendrogram.

CANOCO 4.51 was used to carry out correspondence

analyses for diatoms (Fig. 5a, c) with only diatoms that

formed C5% of the flora in at least one sample (60)

being shown in Fig 5a.

The plotting program C2 was used to construct

stratigraphic diagrams. Environmental reconstructions

for pH were based on training sets and transfer

functions provided by the European Diatom Database

(EDDI, http://craticula.ncl.ac.uk/Eddi/jsp/index.jsp).

This study made use of the combined pH training set

and Locally-weighted Weighted Averaging (LWWA),

which generates a local training set for each fossil

sample based on the 50 closest analogues defined by

the minimum chi-squared distance. Extinct taxa

ranged between about 0–9% of the species present

and were excluded prior to analysis, reducing the

reliability of the results accordingly. Water depths

were assessed qualitatively using known habitat

preferences (Gasse 1986; Ognjanova-Rumenova and

Popova 1996).

Results

Lithofacies

The sediments were examined in five logged sections

(Fig. 2) in the Florina and Kozani Basins and in

several small supplementary outcrops. The main

lithologies include organic-rich lignitic horizons and

clayey siltstones, a variety of diatomaceous and non-

diatomaceous siltstones, sandstones, conglomerates,

diatomites, tufas, marlstones and limestones, as well

as deposits of huntite and magnesite.

One long section was examined in the Florina

Basin at a lignite quarry near Klidi village (Fig. 1a).

Clastic and biogenic materials dominate (Fig. 3a)

with lignites occurring at the bottom of section FK1

(Fig. 2a) and extending for several tens of meters

below the sampled sequence (Fig. 3b). Within the

sampled section (Units 1–5, FK1a, Fig 2a), these

organic deposits consist of alternating black lignite

and yellowish non-diatomaceous siltstones (Fig. 3c).

Fresh horizontally oriented, brown, wood stems and

branches occur locally and there is a lack of evidence

for in situ growth. Individual lignitic beds are

laterally extensive but vary in thickness, having a

wavy form. Many of the coals are rich in gymno-

sperms (Papanicolaou et al. 2000).

White diatomaceous siltstones constitute Units 6–9,

with varying clay content and the deposits being fissile

in Units 7–9. Unit 10 contains organic-rich fissile

siltstones, with common ostracods. Large fragments of

wood lie parallel to the bedding with organics being

more abundant at the base of the sequence, which

laterally grades into lignite.

Units 11–18 are dominated by white to grey

diatomaceous siltstones that show a distinct cyclicity

that has been used to define the seven units in this

part of the succession. They typically consist of a

thick (20–90 cm), massive, white (when dry), diato-

maceous siltstone that passes upwards into a thin

(2–10 cm) grey clayey siltstone (Fig. 3d) along

which the different units tend to break. Rare pebbles

(\3 cm) occur in Unit 14. Whole and fragmented

leaves of angiosperms and various aquatic macro-

phytes are scattered throughout these sediments

(Fig. 3g–j) and are especially abundant in Units 14

and 18 where they are associated with the develop-

ment of vivianite (Fe3(PO4)2�8(H2O)) (Fig. 3f).

Units 19–28 consist of generally thicker (about

70–600 cm) sequences of massive white diatoma-

ceous siltstone. Leaves are less common in Units 19–

21, with minor vivianite in Unit 19. The succeeding

Units 24–28 contain abundant leaf fossils with one

flower fragment (Fig. 3k) in the base of Unit 26. A

distinct layer rich in anapaite (Ca2Fe (PO4)2�4(H2O))

nodules (\1-cm diameter) forms Unit 29 (Plate 1e).

The nodules (faecal pellets?) extend into the base of

Unit 30, which is a massive, 260-cm-thick, grey

diatomaceous siltstone. Anapaite nodules also occur

in Unit 31, which consists of alternating wispy white

diatomite and grey clayey diatomaceous siltstone.

Grey siltstones dominate in Units 32–36, except for

Unit 33, which is lighter in color and Unit 35, which

is comprised of alternating clayey diatomaceous

siltstones and diatomites.

The sediments in the Kozani Basin were sampled

at several locations and consist of a variety of clastic,

346 J Paleolimnol (2010) 44:343–359

123

biogenic and chemical sediments (Fig. 4). The rock

types of the Lower Neogene Series at Kariditsa

(Figs. 1b, 2b, 4a) are comprised mainly of fine-grained

lithologies (claystones, siltstones, sandy siltstones)

with many horizons containing gastropods. A promi-

nent lignite constitutes Unit 11 in Section KK1

(Fig. 2b). The deposits of the Upper Neogene Series

(KP1, KM1 and KM2, Figs. 2b, 4b–f) are more varied.

They include siliciclastic claystones, siltstones, sand-

stones and conglomerates, with ripple cross-laminae

and cross beds in several sandstones. Locally, sand-

filled channels are present. Gastropods occur in both

siliciclastic and biogenic (diatomaceous siltstone,

diatomite, tufa, limestone) facies. Chemical sediments

are dominated by huntite (Mg3Ca(CO3)4), magnesite

(MgCO3) and hydromagnesite Mg5(CO3)4(OH)2�4(H2O)

that were deposited in small shallow late Neogene

alkaline lakes (Wetzenstein 1975; Stamatakis 1995).

The diatom floras

A total of 133 taxa was recognized in the sections

from the Florina and Kozani Basins. Extinct species

accounted for 12% of the total flora, but locally

accounted for 0–31% of the diatoms in the assem-

blages. Preservation ranged from poor to excellent

with corrosion being more prevalent in association

with the Upper Neogene chemical sediments of the

Kozani Basin. Figure 5b shows a cluster analysis of

the most common species observed in all of the

sampling sites. Six major groups can be distinguished

at the 0.42 level of dissimilarity, which corresponds

with a major step in the clustering.

Group 1 is comprised of Aulacoseira ambigua

(Grun.) Simons., which forms near-monospecific

floras (up to 99.5%) in diatomites or highly diato-

maceous deposits (Units 30–33 of FK1, Fig. 2)

* **

*

*

1.5 m gap

2 m gap

2m gap

Section FK1

4

12

1

35

6

7

89 10

11131415161718192021

2

Lower Neogene

UpperNeogene

KP1

KK1

KM2KM1

nisabinazoKbnisabanirolFa

3 m

etre

s

White siltstoneLight grey siltstone

Clayey siltstone

Yellowish siltstone

Conglomerate

Sandy siltstone

Massive sandstone

Thin bedsFissile white siltstoneCherty limestoneClaystoneCross-bedded ss.

Limestone

Huntite/

MagnesiteHuntiteSiliciclastic

Brown siltstone

Lignite and wood1

2 3 4

5 6

7

8 9 10

11 12 13

14

15

16

17

18

19

21

20

23

22

24

25

26

27

28

29

30

31

32

33

34

35

36

37

1 2 3 4 5 6 7 8 9

10

11

12

13 14 15 16 17

18

19

20

1

2

3

1

2

3

4

5

6

7

8

9 10

11

12

13 14 15 16 17 18

Diatom samples

Stratigraphicunit

FK1a

FK1b

FK1c

FK1d

Pyrite noduleLeavesFlower

*

*

* VivianiteAnapaite

OstracodsGastropods

hydromagnesite/magnesite

Peb

bles

Fig. 2 Lithological logs.

a Klidi Quarry section,

Florina Basin. b Kozani

Basin sections. Locations

are shown in Fig. 1

J Paleolimnol (2010) 44:343–359 347

123

towards the top of the Klidi Quarry section (Florina

Basin). Occasional Staurosira construens Ehr. are

also present, but in low numbers.

Group 2 floras occur in diatomaceous siltstones in

several locations in both the Kozani and Florina

Basins. The group is characterized by a greater

Fig. 3 Sediments at the Klidi Quarry, Florina Basin. a General

view with white to grey diatomaceous siltstones overlying

darker lignites; b Alternating lignites and siltstones, white

bar = 1 m; c Detail of the basal part of the Klidi section

showing the transition from lignites to diatomaceous sediments

(Unit 6 upwards), tape = 1 m; d Cyclic diatomaceous sedi-

ments with thin and darker clayey siltstones at the top of each

unit, tape = 1 m; e Anapaite nodules (black arrow) in clayey

diatomaceous siltstone; f Vivianite alteration (black arrows) of

leaf debris; g–j Well-preserved leaf fossils; k Flower present in

Unit 26; l Conglomerates from the top of the succession, which

rest unconformably on the diatomaceous deposits

348 J Paleolimnol (2010) 44:343–359

123

variety of species than Group 1 and could perhaps be

further subdivided (Fig. 5b) with S. construens and

Pseudostaurosira brevistriata (Grun. in V. Heurck.)

Will. & Round forming one sub-group (2A, Fig. 5b).

Subgroup 2B is comprised of Actinocyclus makaro-

vae Temn. & Ognjan., Ellerbeckia kochii (Pant.)

Temn., Val. & Ognjan., Cyclotella castracanei Eul.,

C. iris, Cyclotella iris var. ovalis Brun & Herib.,

Fragilaria bituminosa Pant., Pseudostaurosira brev-

istriata (Grun. in V. Heurck) Will. & Round and

Pseudostaurosira zeilleri (Herib.) Will. & Round.

The third major grouping includes two unidenti-

fied Cyclotella species, plus Cyclotella iris var.

cocconeiformis Brun & Herib., Cyclotella iris var.

integra Perag. & Herib.. Aulacoseira distans (Kutz.)

Simons. also falls within this group, but with a

relatively higher level of dissimilarity separating it

from the Cyclotella taxa (Fig. 5b). This flora was

confined to diatomaceous siltstones in the Klidi

Quarry. Group 4 is also dominated by Cyclotella

species, which are dominated by C. elymaea and

Cyclotella ocellata Pant. Group 4 occurred in highly

diatomaceous siltstones in the upper part (above Unit

27 in FK1, Fig. 2) of the Klidi Quarry section in the

Florina Basin.

Group 5 is comprised of Anomoeoneis sphaero-

phora (Kutz.) Pfitz., which generally constitutes close

to 100% of the flora, but with the occasional presence

of an unidentified Navicula designated ‘‘sp. 5’’. This

grouping was present in the upper parts of the lower

Neogene Kariditsa section (KK1), where it was found

in claystones.

Group 6 is the most diverse of the clusters that

were recognized and is comprised of a variety of

Fig. 4 Sediments from the Kozani Basin. Black vertical bars

are approximately 2 m. a: Kariditsa. Diatomaceous and non-

diatomaceous siltstones and claystones with a dark lignite layer

towards the base; b: Huntite/hydromagnesite quarry section

(KM2; Fig. 2) near Neraida, diatoms absent from the sequence;

c: Magnesite Quarry section (KM1; Fig. 2) near Neraida,

diatoms rare in the white basal magnesite and abundant in the

overlying siltstones and a carbonate that lies between these two

major units; d: Tufa mound at the huntite/hydromagnesite

quarries near Neraida village, diatoms rare; e: Section KR1

near Neraida. Diatoms are absent except in the upper 4 m. f:

Section KP1 at Stena Portas (old magnesite quarry). White

magnesite-rich beds at the base gives way to about 2 m of

sediment with rare to moderately abundant diatoms. Diatoms

absent in the upper sediments

J Paleolimnol (2010) 44:343–359 349

123

benthic and epiphytic diatoms that are largely con-

fined to the Kozani Basin. The flora can perhaps be

further subdivided at a low dissimilarity level of 0.08

(Fig. 5). Subgroup 6a (Cymbella helvetica Kutz.,

Epithemia sorex Kutz., Staurosirella leptostauron

(Ehr.) Will. & Round) occurred in siltstones at

Kariditsa (KK1, Fig. 2). Subgroup 6b (Cyclotella

aegeae Ec,-Amilli, Navicula bituminosa Pant.) was

confined to the base of the Stena Portas sequence

(Units 2–10 in KP1, Fig. 2), where it formed a sparse

flora in siltstones associated with reworked huntite.

Diatoms belonging to Subgroup 6c (Cymbella, Gom-

phonema and Eunotia floras, plus Nitzschia sinuata

var. tabellaria Grun.) occur mainly at Kariditsa

(KK1, Figs. 1 and 2) in siltstones, often with

gastropods. Group 6d includes a diverse range of

diatoms (e.g., Synedra ulna (Nitz.) Ehr., Fragilaria

hungarica Pant., Nitzschia frustulum (Kutz.) Grun.;

a b

c

d

Fig. 5 Multivariate

Analyses of the diatom

data. a Correspondence

Analysis for diatom species;

b Cluster analysis for

diatom species; cCorrespondence Analysis

for samples from the Klidi

and Kozani areas; d Cluster

Analysis for samples

350 J Paleolimnol (2010) 44:343–359

123

Fig. 5b), which also occurred present in siltstones at

Kariditsa.

Cluster analyses show a clear separation of the

samples from the Florina and Kozani Basins, with the

exception of a small number of outliers. For example,

sample R06 from the Kozani area appears to be related

to samples from the middle and lower parts of the Klidi

section in the Florina Basin. The general spatial

separation of samples is clearly visible in correspon-

dence analyses (CA) (Fig. 5c) with Kozani samples

plotting in Quadrant IV and part of Quadrant II. The CA

for the Klidi Quarry material also shows a distinct trend,

with samples from the basal part of the sequence lying in

Quadrant III and those from the top of the deposits

occurring in Quadrant II. Figure 5a presents a CA for

species, with diatoms belonging to the six major groups

identified from cluster analysis shaded. These data

reinforce the separation of these various groupings.

Diatom stratigraphy

Detailed diatom stratigraphies were generated for two

of the longest sections. Figure 6 shows the results for

Section FK1 from the Klidi Quarry. The sediments are

mainly dominated by Cyclotella, A. ambigua and

several species of Fragilariaceae, with S. construens,

P. brevistriata and Staurosirella pinnata (Ehr.) Will. &

Round occurring to varying degrees throughout the

diatomaceous parts of the sequence. Ten major stages

(A–J) can be recognized based on the diatom floras and

diatom abundance in the sediments. Only rare and

fragmentary diatoms occur in Stage A deposits, which

consist of alternating lignites and pale yellowish

siltstones. Epithemia and Cymbella taxa characterize

Stage B, together with A. ambigua and Cocconeis

placentula Ehr.. Cyclotella sp. 1 is similar to a

cyclotelloid taxon, with the same designation,

described by Ognjanova-Rumenova (2005) from the

Bitola Basin. She notes its resemblance to C. ocellata,

but with differences in terms of marginal and central

fultoportulae, the presence of spines and a variable

elliptical to circular shape. Other common taxa include

Fragilaria bituminosa Pant, P. brevistriata, S. constru-

ens and S. pinnata.

Stage C deposits are comprised of diatomaceous

siltstones and diatomites and are characterized by the

presence of C. elymaea and relatively abundant

C. iris var. ovalis. Other common taxa include

Cyclotella sp. 1, F. bituminosa, P. brevistriata,

S. construens and S. pinnata. Cymbella and Epith-

emia taxa disappear at the base of this stage, which

can perhaps be subdivided further based on the higher

percentages of S. construens and S. pinnata in

Substage Ci (Fig. 6).

Cyclotella elymaea disappears in the diatomites

that comprise Stage D, which can be subdivided

using C. iris var. ovalis. This taxon is less common

than S. construens in Substage Di, but more abundant

in Substage Dii. Cyclotella elymaea is again present

in Stage E sediments, with C. iris becoming a

common diatom and C. iris var. ovalis remaining a

major part of the flora.

Cyclotella castracanei and C. ocellata form impor-

tant elements in the flora for the first time from the basal

diatomites of Stage F. Cyclotella elymaea remains

common while C. iris declines and C. iris var. ovalis

remains an important element of the flora. Cyclotella

sp. 1 occurs only at trace levels of \1% in the

diatomaceous silts of Stage G, and C. ocellata becomes

dominant. Cyclotella castracanei remains common.

Diatomaceous silts and diatomites constitute Stage

H, which is distinguished by the abundance of

A. ambigua. At several horizons this species forms

nearly monospecific floras in diatomite. Cyclotella

elymaea disappears from the flora as do Actinocyclus

makarovae and E. kochii, which form a small (\5%)

component of the floras throughout Stages D–G and

parts of Stage B and C. Diatoms are absent or occur

as rare fragments in the siltstones and claystones of

Stage I, but reappear in Stage J where C. castracanei

is co-dominant with C. ocellata.

The second diatom stratigraphy (KK1, Fig. 7) was

developed for outcrops at Kariditsa and represents the

lower Neogene of the Kozani Basin. Nine stages can be

identified. Stage A occurs in white siltstones domi-

nated by Navicula bituminosa. The overlying, gastro-

pod-rich, white siltstones contain a sparse (Stage B)

flora dominated by Navicula sp. 4, Neidium affine

(Ehr.) Pfitz. and Neidium iridis (Ehr.) Cl.. Diatoms are

more common in the Stage C siltstones, which also

contain common gastropods and a diatom flora dom-

inated by N. bituminosa with less common Synedra

ulna (Nitz.) Ehr., and several taxa belonging to

Gomphonema, Eunotia and Cymbella (Fig. 7). Stage

D is confined to a relatively thin clayey siltstone and is

dominated by S. leptostauron.

Diatoms are absent in the fissile, gastropod-rich,

white siltstones, cherty limestone, claystone and

J Paleolimnol (2010) 44:343–359 351

123

lignite that make up Stage E. The succeeding Stage F

is dominated by A. ambigua in siltstones with a few

leaf fossils. S. ulna and Eunotia spp. are common as

is Navicula sp. 5. Stage G occurs in similar

sediments, but is dominated by S. ulna, A. sphaero-

phora and several Gomphonema and Cymbella taxa

plus Nitzschia sinuata var. tabulata. Diatoms are

again absent in Stage H, which occurs in siltstones

and claystones with gastropods. The top of the

sequence comprises Stage I where diatoms are sparse

and comprised solely of A. sphaerophora.

The diatoms in the Upper Neogene Series of

Kozani were detected in a small outcrop, close to the

old Aliakmon Bridge (KS1; Fig. 1) and in parts of

several longer sections (KR1, KM1, KM2, KP1;

Figs. 1 and 2). A sparse diatom flora occurred in the

lower parts of the sediments at Stena Portas (Fig. 1).

There, Units 1 and 2 consist of huntite and magnesite

and lacked diatoms (KP1, Fig 2). Units 2–6 and 8

and 9 contained a sparse flora dominated by

C. aegeae, C. elymaea and N. frustulum, together

with A. sphaerophora and Thalassiosira sp. 1. Unit 7

contains moderately common diatoms and is domi-

nated by Cyclotella sp. 3. Units 10 and 11 are domi-

nated by moderately common C. aegeae (81–85%) and

Cyclotella sp. 3. Diatoms are rare and fragmentary in

the remaining Units 12–20.

The huntite/hydromagnesite quarries of Neraida

that belong to the Upper Neogene Series were exam-

ined at sections KM1 and KM2 (Figs. 1 and 2).

Diatoms were absent or scarce in KM2, but were

present in all units of KM1. In the latter section, Unit 1

is comprised of huntite/hydromagnesite with diatoms

forming\5% of the deposit. The most common taxa

40 0

400

800

1200

1600

2000

2400

2800

3200

3600

100%

Abu

ndan

ce

50%

Cyc

lote

lla c

astr

acan

eiC

. dis

tingu

ida

C. e

lym

aea

C. i

risC

. iris

v. c

occo

neifo

rmis

C. i

ris v

. int

egra

C. i

ris v

. ova

lisC

. oce

llata

Cyc

lote

lla s

p. 1

Cyc

lote

lla s

p. 2

Act

inoc

yclu

s m

akar

ovae

Elle

rbec

kia

koch

iiA

ulac

osei

ra a

mbi

gua

A. d

ista

nsA

. gra

nula

taF

ragi

laria

bitu

min

osa

Pse

udos

taur

osira

bre

vist

riata

Sta

uros

ira c

onst

ruen

s

Fra

gila

ria h

unga

rica

Sta

uros

irella

lept

osta

uron

Sta

uros

irella

pin

nata

Pse

udos

taur

osira

zei

lleri

Coc

cone

ois

plac

entu

la

Nav

icul

a sp

p.A

chna

nthe

s sp

p.C

ymbe

lla s

pp.

Epi

them

ia s

pp.

Chy

soph

ycea

n cy

sts

Planktonic Facultative planktonic Benthonic Epiphytic

7 8 950pH

Habitat pH

Habitats:

Stage

A B Ci

Cii

Di

E

F

G

H

Diatoms rare, broken or absent Diatoms rare, broken or absent

tnesba ro nekorb ,erar smotaiDtnesba ro nekorb ,erar smotaiD

FK1

IJ

Dii

20

22

26

27

28

30

19

14

18 17

21

32 34

36

23

6

0 20 % diatomsand cycts % %

cm

Fig. 6 Diatom and chrysophycean stomatocyst stratigraphy for FK1, Florina Basin. Habitat and pH data are shown to the right.

Location is shown in Fig. 1

352 J Paleolimnol (2010) 44:343–359

123

included E. sorex (16%), S. pinnata (14%), Cymbella

cistula (Ehr.. in Hempr. & Ehr.) Kirchner (10%),

P. brevistriata (8%) and N. affine (6%). Unit 2 is a

carbonate dominated by E. sorex (22%), Navicula sp. 7

(20%) and C. helvetica (9%), N. frustulum (8%) and

C. ehrenbergii Kutz. (7%). Unit 3 consists of diato-

maceous brown siltstones with abundant and varied

diatom floras that include: C. ocellata (4–48%),

C. helvetica (20–42%), P. brevistriata (1–16%),

E. sorex (6–9%), S. pinnata (1–8%) and Gomphonema

parvulum (Kutz.) Kutz. (0–6%). Spring tufa mounds

within the huntite/hydromagnesite quarry floor yielded

no diatoms although a nearby stromatolitic carbonate

contained occasional S. leptostauron (20%), C. helve-

tica (20%) C. cistula (8%), A. granulata (8%) and

Mastogloia elliptica (Ag.) Cl. (5%).

Discussion

Environmental conditions in the Klidi area

Previous studies (Ehlers 1960; Stamatakis 1994,

1995; Steenbrink et al. 2006) have established the

existence of Late Miocene to late Early Pliocene

sedimentation in the Florina-Ptolomeis-Kozani chain

of basins. Several tens of metres of alternating

lignites and siltstones underlie the stratigraphic

section reported in this study. However, Oikono-

moupolos et al. (2008) have carried out detailed

investigations of these basal lignites at Achlada,

about 10 km north of Klidi. There, the organic

deposits have been reported as forming in peat

swamps on a floodplain crossed by a major mean-

dering river system, with at least some of the lignite

forming in an oxbow lake setting (Fig. 8a).

Alternating lignite seams and siltstones occur at

the base of FK1 (Fig. 2) and form the top of the

succession. Transported organics (wood fragments)

are present in the siltstones, but there is a lack of

rooted plants suggesting an origin by reworking of

swamp materials as a transgressing lake was inun-

dating them. Diatoms became well established with

the laying down of Unit 6 (Fig. 2) and the develop-

ment of Stage B floras (Fig. 6). These contain a

mixture of planktonic and facultative planktonic

diatoms and have the highest proportion of epiphytes

in the sequence. Transfer-function data (Fig. 6)

pH

Habitat pH

50%

Abun

danc

e

50%

Aulaco

seira

ambigu

a

Frag

ilaria

bitu

minos

a

Stau

rosira con

struen

s

Stau

rosirella

leptos

tauron

S. pinna

ta

Syne

dra ulna

Achn

anthidium m

inutissim

um

Coc

cone

is place

ntula

Anom

oeon

eis sp

haerop

hora

Nav

icula am

phice

phala

N. b

ituminos

a

Nav

icula sp

. 4

Nav

icula sp

. 5

Neidium

affine

N. Irid

is

Gom

phon

ema ac

uminatum

G. intric

atum

G. p

arvu

lum

Gom

phon

ema sp

. 2

Epith

emia sorex

Euno

tia pec

tinalis

E. lu

naris

E. prearup

ta

Amph

ora lib

yca

Cym

bella

asp

era

C. c

istula

C. turgida

Nitz

schia sinu

ata v. ta

bulata

6 8 100 50% 50%

Stage

AB

C

E

FG

H

I

D

KK1

0

50

100

150

200

250

300

350

400

450

500

550 cm

4

12

1

356

7

89 10111314151617181920

21

2

Stra

tigra

phic

units

Planktonic Facultative planktonic Benthonic Epiphytic Habitats:

Diatoms rare, broken or absent

Diatoms rare, broken or absent

Fig. 7 Diatoms stratigraphy for Section KK1, near Kariditsa, Kozani Basins. Location is shown in Fig. 1

J Paleolimnol (2010) 44:343–359 353

123

suggest freshwater conditions with pH values of

about 7.2–8.0.

Stage C is characterized by an overall increase in

planktonic diatoms. In Stage Ci, epiphytes decline

from their Stage B percentages and planktonic

species form their lowest contribution to the flora in

the entire section, suggesting very shallow open

water conditions. Transfer-function data indicate a

pH of up to 8.2. Subsequently (Stage Cii), there was

an increase in planktonic C. iris var. ovalis—an

oligotrophic diatom favoured by carbonate-rich water

and a pH of about 7.4 (Serieyssol 1984). Within each

of the depositional units (Units 11–18, FK1, Fig. 2)

of Stage Cii there is a gradual increase in planktonic

diatoms (e.g. C. iris var. ovalis) upwards followed by

a return to floras that suggest shallow water. The

sediments also document a distinct cyclicity with

each unit comprising a highly diatomaceous

sequence, at decimeter to metre thicknesses, passing

upwards into a thin (2–5-cm) clayey siltstone topped

by a sharp bedding plane. Cyclic sedimentation has

been previously reported from several other Florina-

Ptolemais-Kozani Basin (van Vugt et al. 1998;

Steenbrink et al. 1999, 2006) where cycles at

differing scales have been related to precession and

eccentricity orbital periods.

Stage C is also distinguished by a decrease in

diatom abundance from about 95% down to about

10% of the sediment (Units 16–17; Fig. 6), perhaps

related to increased river inputs and dilution of the

diatoms by clastic materials. This decline in diatom

abundance is also associated with the occurrence of

an increase in leaf fossils (Fig. 2), which are locally

abundant and associated with the development of

vivianite. Stamatakis (2004) notes similar leaf-vivia-

nite associations in clayey diatomites from other parts

of the Florina-Ptolemais-Kozani Basin and attributed

their origins to phosphorous enrichment caused by

high biological productivity in the catchment, with

additional phosphorous supplied by the decay of

leaves. Organic decay, or iron-reducing bacteria,

would have assisted in the reduction of iron to the

ferrous form, perhaps under a stratified water body

with anoxic bottom waters, or within deeper anoxic

muds. This would also be consistent with the

generally excellent preservation of leaves. The ideal

pH conditions for vivianite formation range from 6.4

to 7.8 (Nriagu and Dell 1974), values that are

Xi

Xiii

Xii

XivYi

Yiii

Yii

Yiv

X

Y

Meandering river Oxbow lakes

peat

peat

peat

Peat coverted to lignite after burial and time

Shallow swampy lake

Moderately deep lake

Lignite andsiltstone

Diatomaceoussiltstone

Shallowphase

Deepphase

a c

b Pleistocene fluvial pebbles Lake cycles

Miocene lignites

Saline pondsLake

Tufa moundsFloodplain withToxidaceae andmixed forest

(i)

(ii)

Fig. 8 Depositional

models for the Klidi Quarry

area, a–b Florina Basin.

‘‘a’’ represents basal

Miocene lignite

paleoenvironment; ‘‘b’’

shows lake cyclicity for the

main diatomaceous

siltstones and diatomites.

Low lake stages (i)

repeatedly expanded (ii)

and then contracted again.

These variations occurred at

two different time scales

involving multiple shorter

cycles (Xi–iv, Yi–iv) nested

within longer-term changes

(X, Y). c Sedimentation

model for the upper

Neogene of the Kozani

Basin. Tufa-forming springs

and local streams fed saline

ponds in which magnesium-

rich carbonates were laid

down. During low lake level

stages these deposits were

reworked into adjacent

deltaic settings (after Calvo

et al. 1995)

354 J Paleolimnol (2010) 44:343–359

123

consistent with the diatom data that show lacustrine

pH values falling to about 7.5 in the sediments that

contain vivianite.

The succeeding sedimentary units lack the distinc-

tive clayey siltstones at the top of each diatomaceous

siltstone sequence and contain fewer leaf fossils than

the deposits of Stage C. Diatoms are relatively

abundant and there is evidence for continued diatom

cyclicity. Overall, planktonic taxa reach their peak at

the top of Stage D (Dii = top of Unit 21) with a flora

dominated by C. iris var. ovalis. In contrast, substage

Di is dominated by S. construens, an alkaliphilic

epiphyte capable of living in the plankton of shallow

lakes (Gasse 1986). Although this latter taxon

suggests shallow conditions, proceeding upwards in

the profile, planktonic diatoms are also present and

display an percentage increase within Units 19 and 21,

although there is an decline through Unit 20. Data for

pH suggests consistent values of about 7.6 for

substage Di, falling to about 7.1 in Dii.

In Stage E there are again peaks in planktonic

diatom percentages at the top of each of the

sedimentary units (22–27) although the floras are

now also characterized by the presence of C. iris and

C. elymaea. Cyclotella iris occurs in the bottom mud

or phytoplankton of alkaline lakes (Gasse 1986). The

ecological requirements of the C. elymaea are

uncertain given its extinct status, although Ecomo-

mou-Amilli (1991) suggested an association with

‘‘higher salinities and alkalinities’’ than were typical

of the Florina-Ptolemais-Kozani Basin lakes. Trans-

fer-function data for extant species suggest a pH of

7.3–7.7. Diatom abundances decline slightly, leaf

fossils (and one flower, Fig. 3k) are again common,

and vivianite occurs at several horizons—features

that resemble those of Stage C, but with a less drastic

decline in diatom abundance.

Cyclotella castracanei appears for the first time in

Stage F, which again shows an upward increase in

planktonic floras through Unit 28. The pH data

indicates a decline from about 7.7 to 7.2. This inverse

relationship between planktonic diatoms and pH is

repeated at many levels through the sequence

(Fig. 6). A 3-cm-thick layer of diatomaceous clayey

silt with abundant euhedral bladed anapaite crystals

with a greenish color that form nodules (Fig. 3e)

comprises Unit 29 at the top of Stage F. Stamatakis

(2004) suggested that these nodules (up to 1-cm

diameter) formed diagenetically at the expense of

vivianite when exposed to Ca2? and HCO3--rich

meteoric waters and/or somewhat acidic groundwa-

ter. This level in the sequence is also associated with

a moderate drop in lake water pH according to the

diatom record.

Light grey siltstones dominate through the remain-

der of the section with occasional pure diatomites

occurring in Units 31 and 33. The flora in Stage

G (lower Unit 30; Fig. 6) shows a shift towards

C. ocellata, a freshwater species that occurs in

shallow water bodies. Transfer-function data indicate

a pH of about 8 to 8.4. Gasse (1986) suggests it is

favored by oligotrophic conditions. Stage G does not

appear to document any increase in the percentage of

planktonic taxa with height through the section.

However, Stage H does display an increase in plank-

tonic diatoms, as well as a switch from C. ocellata

to A. ambigua. This planktonic species is favored

by relatively shallow water and low alkalinity

(\5 meq l-1) (Owen and Utha-aroon 1999) with

derived pH values of about 7.5–8.

Stage I contain only rare and broken diatoms in a

light grey siltstone (Unit 34) and in streaky white

siltstones in a grey siltstone (Unit 35). Anapaite

nodules are scattered throughout this latter deposit,

which suggests post depositional alteration of a

vivianite precursor. Collectively, the data document

a possible period of emergence prior to renewal of

aquatic conditions in Stage J. A period of erosion

followed with later deposition of fluvial conglomer-

ates (Fig. 2).

Further data can be derived from chysophycean

stomatocysts, which are common at several levels in

the sedimentary sequence, forming 0–20% of the

total diatom ? cyst assemblage (Fig. 6). These

organisms are most abundant in low productivity

lakes with circum-neutral pH, low alkalinity and low

phosphorous (Ognjanova-Rumenova 2005). Smol

(1985) has used the ratio of diatoms to cysts as an

indicator of trophic state. The data for Klidi (Fig. 6)

show that cysts are most common in the lowest part

of the succession (Stage Cii) in sediments with low

diatom abundance. The ratios thus document a period

of oligotrophic conditions. Cysts also tend to be

absent or form a low contribution during periods

when planktonic diatoms were most common perhaps

reflecting periods of increased trophic state.

In general, there appear to be multiple episodes of

diatom cyclicity at varying scales (Fig. 8b). Several

J Paleolimnol (2010) 44:343–359 355

123

diatom stages (C, D, F and H) show an increase in

planktonic diatoms proceeding upwards in the profile.

At a smaller scale, many individual sedimentary units

also tend to show increases in planktonic species. The

floras of the Klidi Quarry are similar to those reported

from Bitola (Fig. 1) in Macedonia (Ognjanova-

Rumenova 2005) where the floras are also dominated

by Cyclotella spp., with common S. construens.

However, no cyclic patterns are reported from that

study, which might reflect variations in local condi-

tions or, perhaps more likely, the wider sampling

interval (17 samples from about 63 m of core), which

produced a less detailed diatom record. In the same

study, Ognjanova-Rumenova (2005) also pointed out

that the appearance of C. castracanei and disappear-

ance of Miocene Actinocyclus marked the lower part

of the Pliocene, which in the section reported here

occurs in Stages F–G.

Environmental conditions in the Kozani Basin

The lower Neogene of the Kozani Basin is

represented in our sampling set by the Kariditsa

sedimentary sequence (Figs. 2 and 7). These depos-

its are a complex mix of claystones, siltstones, and

minor lignite. Gastropods are present in many

horizons. Diatoms are absent from about 50% of

the sequence and occur at three major levels.

Benthonic floras that give way to an increasing

number of epiphytes dominate Stages A–D. The pH

ranged between about 6.8 and 7.8, somewhat lower

than pH data for the Klidi area. Diatoms are absent

in Stage E and reappear with a mixed benthonic-

epiphytic flora and common Eunotia spp., which

indicate even lower pH values (6.6–7.2). Diatoms

are absent in Stage H, with a sparse flora domi-

nated (up to 100%) by A. sphaerophora character-

izing Stage I. This taxon is usually cited as an

indicator of shallow water bodies with a medium to

high alkalinity ([10 meq l-1). It also occurs in

hyperalkaline settings.

In general, the data indicate that the Kariditsa

sediments formed under very shallow acidic to

neutral water, with several periods of desiccation

and emergence occurring. These environments would

also have been favorable to the development of peaty

swamps and the formation of the lignite parts of the

succession. Towards the end of the depositional

period there was a shift towards more alkaline

conditions, perhaps reflecting evaporative concentra-

tion and/or a nearby spring water source.

The upper Neogene sequences contain evidence of

both high and moderate salinity conditions. Calvo

et al. (1995) have noted the occurrence of magnesite

along with huntite, hydromagnesite and dolomite at

several locations in the Kozani Basin. Outcrops on

the southern (KP1, Stena Portas) and eastern (KM1

and KM2, Neraida; Fig. 1) margins of the area

contain extensive deposits of primary and reworked

huntite and magnesite. These appear to have formed

in small spring-fed saline lakes on the margins of a

larger, fresher water body (Fig. 8c). The Neraida

outcrops, for example, contain extensive beds of fine-

grained snow-white huntite/magnesite with minor

aragonite and magnesite, associated aragonitic tufa

mounds and local stromatolites. The magnesium-rich

deposits contain a sparse, fragmented diatom flora

dominated by shallow-water epiphytic and benthic

taxa that would be incompatible with a highly saline

environment. These diatoms may therefore have been

transported into the saline lakes from adjacent

swampy areas or formed part of a freshwater river

flora. Epiphytic Cymbella spp. are more abundant and

dominate, together with E. sorex, in the overlying

carbonate and diatomaceous siltstone units (Fig. 2),

documenting shallow-water sedimentation. Diatoms

at these higher levels are also more varied and

abundant suggesting extensive freshwater conditions.

Section KP1 (Figs. 1 and 2) at Stena Portas also

contains magnesium-rich chemical sediments. Calvo

et al. (1995) noted the presence of dolomicrite and

marl that formed in fresh aquatic environments and

attributed the presence of huntite to small saline

lakes, probably similar to those near Neraida.

Reworked huntite was likely eroded from these

ponds and incorporated in deltaic facies entering a

larger water body. Diatoms are generally absent in

the Stena Portas deposits, except near the base of

the sequence where C. aegeae, C. elymaea and

N. frustulum, together with A. sphaerophora and

Thalassiosira sp. 1. The first two of these species

are known from the fossil record and have poorly

understood ecologies, although Ecomomou-Amilli

(1991) noted that C. elymaea was associated with

chemical sediments suggestive of higher salinities.

N. frustulum occurs in a variety of settings, but

A. sphaerophora is suggestive of higher salinities

and alkalinities.

356 J Paleolimnol (2010) 44:343–359

123

Conclusions

The Late Miocene through Pliocene sedimentary

rocks of the Florina-Ptolomeis-Kozani Basin include

lithofacies of biological, chemical, and detrital origin.

These host six distinct diatom groups and several

subgroups in cluster analyses that appear to be related

to the type of sediments in which they are found.

Group 1 was associated with very pure diatomites and

comprised mainly A. ambigua, while highly diato-

maceous deposits tended to be associated with Group

4 diatoms (mainly C. elymaea and C. ocellata).

Group 2 (Cyclotella spp. and Fragilariaceae, and

other taxa) and Group 3 (two unidentified Cyclotella,

plus C. iris var. cocconeiformis, C. iris var. integra,

and A. distans) were present in diatomaceous silt-

stones. Group 5 (A. sphaerophora, Navicula sp. 5)

was associated with claystones. Group 6 is dominated

by varied benthic and epiphytic taxa and was found in

siltstones that were locally rippled and/or also

contained gastropods. Group 6b appeared to be

present where chemical sediments such as huntite

also occurred.

The Klidi section (Florina Basin) consists of

diatomaceous siltstones and diatomites that rest on

a thick sequence of lignites and siltstones. The older

lignitic succession formed on a floodplain with

backswamps. Cyclotella spp. and a variety of Frag-

ilariaceae dominate the stratigraphic section, with

A. ambigua being abundant towards the top of the

sequence. These diatoms record a series of lake

expansions and contractions that occurred on at least

two time scales that may reflect the influence of

precession and eccentricity cycles as noted by

Steenbrink et al. (2006). Biostratigraphic data for

the diatoms suggest that the Mio-Pliocene boundary

lies within diatom stages F and G. Chysophycean

stomatocysts suggest periods of oligotrophic condi-

tions, particularly towards the base of the Klidi

sequence.

The Kozani Basin includes both lower and upper

Neogene deposits. The former were characterized by

siltstones, claystones and lignites with predominantly

epiphytic and benthic diatom floras that document

shallow-water deposition under acidic to neutral pH

conditions. Environments became alkaline at the top

of the succession. The upper Neogene includes

several chemical lithofacies that reflect sedimentation

in small saline lakes. Diatoms and sedimentological

data also indicate the occurrence of local shallow

freshwater environments and springs that probably

discharged alkaline waters into the former lakes.

Acknowledgments The authors thank Alison Lee for her

work on preparing diatom slides. Owen was supported by a

Hong Kong Baptist University Grants (FRG/06-07/I-28 and

FRG/08-09/II-25). Renaut was supported by the Natural

Sciences and Engineering Research Council of Canada,

Discovery Grant 629-2008. Stamatakis received funding from

the University of Athens.

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