In: Diatoms: Ecology and Life Cycle

Editors: James C. Compton

ISBN: 978-1-61761-979-3
2011 Nova Science Publishers, Inc.

Chapter 8

PROBLEMS AND POTENTIALITIES OF USING

DIATOMS AS PALEOCLIMATIC INDICATORS IN
CENTRAL ARGENTINA
Gabriela S. Hassan*, Eleonor Tietze, Claudio G. De Francesco and
Paula A. Cristini
CONICET - Instituto de Geologa de Costas y del Cuaternario, Universidad Nacional de
Mar del Plata. Mar del Plata, Buenos Aires, Argentina.

ABSTRACT
Paleoclimatologists advocate learning from the past to know the natural and
anthropogenically induced climatic shifts that occurred in the history of the Earth, and to
estimate future changes in ecological systems related to the projected range of
temperature increases. Framed within this context, current rates of global warming are
unprecedented based on the last 10,000 years of paleoclimatological evidence. Diatom
analysis has played a major role in the reconstruction of these past climate changes,
bringing evidences of water and air temperature fluctuations, as well as of climatically
driven variations in water chemistry variables. Most of these evidences came from North
American and European records, while the number of studies in South America has been
considerably lower. Hence, many questions regarding the natural climatic cycles and the
strength of human induced changes remain unanswered for vast regions of South
America. In Central Argentina, the occurrence of many shallow lakes with sedimentary
records encompassing the last 10,000 years, as well as outcropping alluvial sequences of
Late Pleistocene and Holocene origin, provide good opportunities for studying these
topics. Moreover, the diverse ranges of temperature and precipitation that characterize
this region would allow studying the patterns of distribution of modern diatom
assemblages against climatically driven environmental variables, as well as to assess the
potential application of these modern datasets to the quantitative reconstruction of past
climates. In this chapter, we analyze the problems and potentialities regarding to the
application of the modern diatom data to assess past climate changes in central Argentina.
The main objective is to identify particular lines of research that need to be addressed in
* Corresponding author: E-mail: ghassan@conicet.gov.ar

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

order to allow precise quantitative reconstructions of Holocene climatic changes in this
region.

INTRODUCTION
The hypothesis that the Earth is currently undergoing anthropogenic global climate
change has accumulated considerable supporting evidence during the last decades (Angeler,
2007). In fact, global mean surface temperature increased by ~0.6C during the 20th century,
and modeling approaches using different CO2 emission scenarios suggested that temperature
will increase another 1.4 to 5.8C by 2100 (IPCC, 2001). These instrumental evidences of
climate warming have lead scientists to question if these tendencies could be a consequence
of natural fluctuations in Earths climatic patterns (Fritz, 1996). In this sense, the
paleoclimatic record brings a natural baseline against which the anthropogenic changes can be
compared, both in magnitude and in rate (Schneider and Mastrandrea, 2005). Framed within
this context, modern rates of global warming appear to be unprecedented when compared
with the natural fluctuations that occurred during the last 10,000 years (Angeler, 2007;
Schneider and Mastrandrea, 2005). This brings new insights into the relative importance of
paleodisciplines such as paleolimnology to provide quantitative information on past climates,
which will have implications for understanding modern deviations from those natural
patterns. The challenge for palaeolimnologists in this framework is to assess how climatically
driven changes are recorded in sediments, to outline techniques for precise climate
reconstruction, and, when possible, to evaluate the impact of climate change on past
ecosystems (Battarbee, 2000).
Diatoms have played a major role in the reconstruction of past climate changes, being
used as proxy indicators to reconstruct Holocene climate variability in every continent
(Mackay et al., 2003). The majority of recent diatom-based paleoclimatological studies apply
quantitative multivariate techniques to reconstruct past environmental variables related to
climate, either directly, such as surface-water temperature (e.g. Vyvermann and Sabbe, 1995;
Rosn et al., 2000; Bigler and Hall, 2002) and air temperature (e.g. Korhola et al., 2000), or
indirectly by reconstructing, for example, salinity (e.g. Fritz et al., 1991; Laird et al., 1996;
Gasse et al., 1997; Verschuren et al., 2000), dissolved organic carbon (DOC; Pientiz et al.,
1999), conductivity (e.g. Davies et al., 2002) and pH (e.g. Psenner and Schmidt, 1992; Koinig
et al., 1998). Studies based on qualitative information provided by assessing changes in
diatom species and assemblages, especially with respect to their habitat, survival strategies
and autoecologies, are also common (e.g. Smol, 1988; Bradbury et al., 2002).
Reconstructions based on all these methods rely on the general assumption that the
environmental requirements of the fossil diatom taxa used as bioindicators have remained
constant during the period considered and, consequently, are similar to those of their closest
living representatives. In this way, the environmental information obtained from living
organisms can be used as modern analogous and extrapolated to the fossil record, particularly
in Quaternary research (Hassan, 2010).
Most diatom-based paleoclimatic records came from North America and Europe, while
the number of studies in South America has been considerably lower (e.g. Gonzlez and
Maidana, 1998; Jenny et al., 2002; Markgraf et al., 2003; Tapia et al., 2003; Hassan et al.,
2004; Moro et al., 2004; Fey et al., 2009). Hence, many questions regarding the natural

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

187

climatic cycles and the strength of human induced changes remain unanswered for vast
regions of South America (Villalba et al., 2009). This is the case of Central Argentina plains,
an area in which the prevailing flat geomorphology promotes the occurrence of numerous
shallow lakes that constitute archives of Holocene environmental changes (Garca-Rodrguez
et al., 2009). Most of these waterbodies are small and shallow, (less than 500 Ha surface and
up to about 4m deep), characterized by conductivities ranging from oligo to hiperhaline,
naturally high nutrient levels in the water column, the absence of riparian forest vegetation,
and the development of dense and rich macrophyte communities (Giorgi et al., 2005). Their
dynamics are intricately linked to climatic conditions and anthropogenic activities (Fernndez
Cirelli and Miretzky 2004; Feijo and Lombardo 2007), as evidenced by the pronounced
hydrological variability that characterized these waterbodies during the 20th century (e.g. lake
level fluctuations, varying river discharges, extension of flooded lowlands; Garca-Rodrguez
et al., 2009). Besides their abundance and climatic sensitivity, very few works intended to
reconstruct Holocene climate changes through the analysis of diatom remains from central
Argentina continental records (Zrate et al., 2000; Hassan et al., 2004; Stutz et al., in press).
Moreover, the potential of modern diatom assemblages as indicators of physical and chemical
parameters of paleoecological significance has been scarcely assessed, and no transfer
functions developed for key environmental variables (Hassan et al., 2009).
We have recently started a project intending to evaluate the relationship between diatoms,
mollusks and environmental variables in the Argentinean Pampa (Hassan et al., 2009; Tietze
and De Francesco, 2010; CREO 01/2010), as well as in the adjacent Espinal (Hassan et al.,
under review), Monte, Patagonian and Andean steppe regions (De Francesco and Hassan,
2009, unpub. data; PICT 2008/0088). As a whole, the three ecoregions cover a wide
environmental gradient, bringing a good opportunity for studying the relationship between
diatom assemblages and physical and chemical parameters under diverse climatological
conditions. It is expected that the resulting data set provide quantitative tools useful for
paleoclimatological research. In this chapter, we analyze the problems and potentialities
regarding the application of diatom-based techniques to assess past climate changes in
Central Argentina on the light of preliminary results of the project. The main objective is to
identify particular lines of research that need to be addressed in order to allow precise
quantitative reconstructions of Holocene climatic changes in this region.

MODERN DIATOM-ENVIRONMENT RELATIONSHIPS IN

CENTRAL ARGENTINA
Although the continental diatom flora of South America is relatively well known
(Frenguelli, 1925, 1942; Rumrich et al., 2000; Metzeltin et al., 2005; Metzeltin and LangeBertalot, 1998), the distribution of taxa in relation to environmental variables has been poorly
documented compared to other regions of the world such as Europe and North America
(Lotter et al., 1997; Rosn et al., 2000; Reavie and Smol, 2001; Bigler and Hall, 2003).
Particularly in Argentina, continental diatoms have been used to infer paleoenvironments in
the southern Pampas (Zrate et al. 2000; Hassan et al. 2004; Stutz et al., in press), the Espinal
region (Gonzlez and Maidana, 1998), and Patagonia (Markgraf et al., 2003; Mayr et al.,
2005; Wille et al., 2007; Fey et al., 2009). However, given the lack of detailed information on

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

local modern diatom distribution in fresh waterbodies, such paleoenvironmental

reconstructions were based mainly on foreign autoecological data. This scarcity of
information has also prevented the development of quantitative models in continental settings
of Argentina.
Diatom species respond to environmental variables directly related to climate, such as air
temperature (Bigler and Hall, 2002; 2003), but are more likely to respond to climate- related
limnological variables (e.g. water depth, salinity, pH, and other water chemistry variables;
Smol et al., 1991). From a paleoclimatic perspective, it does not matter if the relationship is
direct or indirect, only how reliably the variable can be reconstructed and whether the
relationship between climate and indirect factors remains linear over time (Smol and
Cumming, 2000). Hence, the application of diatoms to the reconstruction of past climates
relies on a comprehensive assessment on which environmental variables are related to species
assemblages and to determine how to effectively use these inferences in a paleoclimatic
context (Smol and Cumming, 2000).

Figure 1. Localization of sampling sites.

Up to this moment, a total of 34 lentic environments have been sampled across Central
Argentina (Figure 1). In each survey, samples for diatom, water quality (pH, conductivity,
hardness, total nitrogen, total phosphorous, and silicates) and grain size analyses were
collected (see Hassan et al., 2009 for methodological details). Aquatic vegetation cover at
each sampling site was visually estimated, and a nominal variable was erected for statistical
purposes (0 = no vegetation, 1 = low vegetation cover, and 2 = high vegetation cover; Table
1).The relationship between diatom assemblage composition and the available physical and
chemical variables was explored separately for southern Pampas (Hassan et al., 2009), and
eastern Pampas and Espinal regions (Hassan et al., under review), while data from the Monte,

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

189

Patagonian steppe and Andean steppe regions remains unpublished. Although the dataset is
still small, a preliminary exploration of the general diatom-environment relationship could be
useful to identify general patterns of distribution and to generate new hypothesis for guiding
future research. Hence, in the following sections, results of exploratory analyses based on the
whole data set will be described.
Table 1. Environmental and climatological characteristics of sampled shallow lakes. H:
hardness (mg/l CaCO3), TN: total nitrogen (mg/l), Si: silicates (mg/l); C: conductivity
(mS/cm), V: vegetation cover (0: no vegetation; 1: low; 2: high), ST: maximum summer
temperature (C); MT: mean annual temperature (C), ALT: altitude (m.a.s.l), PRE:
total precipitation (ml/year).
N
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34

Lake name
Latitude
Los Horcones 370331.0S
Salada Grande 365647.6S
Nahuel Ruc 373718.0S
La Brava
375209.0S
Los Pinos
375543.9S
Tupungato
373209.5S
Salada
381553.0S
San Antonio 373753.4S
Blanca Grande 362906.0S
Sauce Grande 385651.6S
Alsina
365251.8S
Encadenadas 380146.9S
Pun
373259.0S
Epecun
370752.9S
El caadn
365130.0S
Lonco May
362330.0S
El Camino
371111.0S
Monte Nievas 355208.0S
La Verde
371516.0S
Las Palomas 361823.0S
Chapalc
365126.0S
Teln
361522.0S
El Bote
364850.0S
La Brava
365544.0S
Horqueta
341019.0S
Presa del Tigre 343628.0S
Presa Nihuil 350121.0S
Diamante
355424.0S
Llancanelo
353903.0S
Blanca
350455.0S
Nia Encant. 350915.0S
Sosneado 2
345043.0S
Sosneado 1
345046.0S
Los Molles
350952.0S

Longitude Ecoregion
H
TN
565801.5W Pampa
687.2
0.5
565849.6W Pampa
743.0
0.5
572539.2W Pampa
255.0
0.5
575825.0W Pampa
96.5
0.5
581934.9W Pampa
173.8
0.5
585553.7W Pampa
173.8
0.5
590820.7W Pampa
124.0
0.5
595214.1W Pampa
147.0 93.6
605310.4W Pampa
227.0 81.3
612050.7W Pampa
558.0 91.3
621322.5W Pampa
267.0 12.8
622841.2W Pampa
263.0 91.5
624627.0W Pampa
241.0 85.5
624826.2W Pampa
1166.0 124.0
633812.0W Pampa
697.0 11.3
635459.0W Pampa
182.0 11.7
641023.0W Espinal 1874.0
3.9
641101.0W Espinal
186.0 23.5
641258.0W Espinal
277.0
8.4
643110.0W Espinal
120.0
2.5
645124.0W Espinal
148.0 12.5
653043.0W Espinal
148.0 22.7
654525.0W Espinal
230.0 34.2
654637.0W Espinal
170.0
5.3
665738.0W Monte
1481.5
0.0
683635.0W Monte
314.0
0.0
684037.0W Monte
630.0
3.7
685315.0W Monte
8391.6
0.0
690949.0W Pat. Steppe 598.0
1.4
693820.0W Andes
1546.6
0.0
695208.0W Andes
337.0 12.8
695501.0W Andes
154.4
0.0
695508.0W Andes
144.4
0.0
695628.0W Andes
323.0
4.9

Si pH
6.5 8.9
3.5 8.9
7.2 8.6
3.6 9.1
22.1 8.2
22.1 8.2
9.8 9.8
12.7 9.0
32.5 8.7
7.5 9.3
2.3 9.0
7.3 9.0
n/d 9.7
1.1 9.2
n/a 10.1
n/a 9.8
n/a 9.8
n/a 10.5
n/a 9.5
n/a 9.7
n/a 10.3
n/a 10.1
n/a 9.5
n/a 10.5
10.0 8.3
12.7 7.7
11.0 8.0
13.2 7.5
5.6 7.9
21.2 8.0
28.5 7.7
32.6 9.0
43.7 9.4
34.5 7.8

C V
5.9 2
5.9 0
1.2 1
0.6 1
1.1 2
1.1 1
5.9 0
4.5 0
0.6 0
6.1 0
1.8 0
1.6 0
7.4 0
99.0 0
3.1 1
7.2 1
15.7 0
1.8 1
1.3 2
0.2 0
8.0 0
3.9 0
0.8 0
0.9 0
9.8 0
0.7 2
1.2 2
30.0 0
8.7 0
3.9 0
0.8 2
0.2 2
0.2 2
0.8 2

ST
27.0
27.2
27.1
27.3
28.4
27.4
28.3
29.6
30.9
28.7
31.6
30.6
31.2
32.0
32.2
32.2
32.2
32.7
32.3
32.4
32.4
33.7
33.3
33.2
33.6
30.0
28.3
28.3
28.4
26.1
22.7
22.0
22.0
23.7

MT
14.7
14.8
14.0
13.6
13.7
13.7
13.7
13.5
15.0
14.5
15.2
14.2
14.4
15.2
15.3
15.5
15.1
15.9
15.1
15.6
15.2
15.8
15.4
15.3
17.0
14.0
12.0
12.1
12.0
10.3
7.3
6.9
6.9
8.2

ALT PRE
2 819
1 784
5 825
88 834
135 831
28 852
83 822
219 782
101 926
11 565
111 806
246 704
237 706
98 715
141 636
152 648
178 504
176 652
178 496
202 632
273 547
307 560
307 504
307 491
416 373
886 291
1321 290
1304 310
1337 233
1654 419
2238 534
2346 624
2346 624
2029 559

Present Climate, Environmental Data and Water Chemistry

The area of interest for the present contribution, located in central Argentina between
parallels 33 and 39 S, comprises five ecoregions: (1) Pampa, (2) Espinal, (3) Monte, (4)
Patagonian Steppe and (5) Southern Andean Steppe (Figure 2; Olson et al., 2001). The first
four regions, which are located along an east to west gradient, show a pattern of decreasing

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

precipitation. In fact, the Pampa region has the highest precipitation values, which vary
between 600 and 1100 mm, whereas in the Patagonian Steppe they vary between 100 mm and
270 mm (Figure 3). This pattern of increasing aridity from east to west responds to the
relative influence of the humid winds coming from the Atlantic Ocean. Temperature varies
between 7 and 17 C across the study area and responds to the latitudinal gradient (Figure 4),
being always higher at the north. Towards the west, the high altitude of the Andes at this
parallel (up to circa 7000 meters above sea level) acts as a barrier for the westerlies from the
Pacific Ocean. This is a mountainous landscape, with deep valleys of glacial origin, and
abundant moraines. The region is located above approximately 2000 m.a.s.l. (Figure 5), and
therefore, its climatic characteristics are mainly related to the high altitude rather than to the
regional precipitation gradient.

Figure 2. Main ecoregions in the study area (according to Olson et al., 2001)

A total of 34 shallow lakes (most < 2m in depth) distributed along the described area
were sampled in this study. Most lakes are thermally homogeneous and saturated with
dissolved oxygen. They are characterized by alkaline (pH 7.34 10.5) and very hard (96.5
8391.6 mg l-1 CaCO3) waters, with muddy sediments (Figure 6). Only those artificial lakes
(reservoirs created in mountainous river valleys) represented towards the west, have hard
substrata, which are mainly dominated by gravel.
Sixteen shallow lakes (sites 1 to 16) were located in the Pampa ecoregion. The sediments
of these lakes are characterized by the highest percentages of organic matter of all the area
(up to 85%; Figure 6). In relation to water chemistry, pH ranges between 8.25 and 9.8, and
hardness between 96.5 and 1166 mg/l CaCO3. Water conductivity is highly variable, between
0.63 and 99 mS cm-1. In agreement with the regional precipitation pattern, both organic
matter content and conductivity decrease towards the west.

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191

Figure 3. Map of annual precipitation of the study area. Data adapted from the WorldClim database
(Hijmans et al., 2005) using the DivaGis 7.1.7.2 program (Hijmans et al., 2009).

Figure 4. Map of mean annual temperatures of the study area. Data adapted from the WorldClim
database (Hijmans et al., 2005) using the DivaGis 7.1.7.2 program (Hijmans et al., 2009).

Eight of the sites (17 to 24) were situated in the Espinal ecoregion, mainly on the
semiarid area of climatic transition between the Pampa and the Patagonian steppe (Paoloni et
al., 2003). The semiarid and arid climates characteristic of this region prevent the
development of an autochthonous hydrographic net. Therefore, it is characterized by many
shallow and saline lakes, usually associated to lowlands without drainage (INTA, 1980). The
chemical composition of the water in these lakes is very variable, and depends on the relief,
climate and composition of the parental material of each hydrological system (Drago and
Quirs, 1996). Conductivities vary between 0.2 and 15.7 mS cm-1, hardness between 120 and
1874 mg l-1 CaCO3, and pH between 9.5 and 10.5.

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

Figure 5. Map of altitudes of the study area. Data adapted from the WorldClim database (Hijmans et
al., 2005) using the DivaGis 7.1.7.2 program (Hijmans et al., 2009).

Figure 6. Grain size and organic matter (LOI) content in sediment samples from 34 sampling sites.

Sites 25 to 28 were located in the semiarid Monte region. These sites include two natural
salt marshes (25, 28) and two artificial reservoirs (26, 27) created on the Diamante and Atuel
rivers, respectively. Consequently, they have contrasting physico-chemical characteristics.
While salt marshes have conductivities between 9.8 and 30 mS cm-1, reservoirs are below 1.3
mS cm-1. Similarly, hardness varies between 1400 and 8400 mg l-1 CaCO3 in salt marshes,
and between 314 and 630 mg l-1 CaCO3 in reservoirs. Water pH was similar, between 7.55
and 8.26.
Only one lake (29) was situated within the Patagonian Steppe region, in southern
Mendoza. This lake, called Llancanelo, is an area of international relevance for biodiversity

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

193

conservation. It was declared to be a Provincial Fauna Reserve in 1980, and was included as a
RAMSAR site in the List of Wetlands of International Importance in 1995 (Ciocco and
Scheibler, 2008). Llancanelo lake is a saline water body (8.68 mS cm-1) situated at a mean
altitude of 1300 m a.s.l. in the Centre-West of Argentina. It has an area of about 65,000 ha
and occupies the depression of the same name generated by compensating sinking during the
rise of the Andes in the Tertiary period. The mean depth of the lake is 30 cm, reaching 2 m on
some sites. The lake is also characterized by alkaline (pH 7.87) and very hard (598 mg l-1
CaCO3) water.
The remaining lakes (sites 30 to 34) were located within the High Andes region. They are
situated at high altitude (between 1337 and 2346 m a.s.l.) and, therefore, subjected to extreme
fluctuations in water temperature. The scarce precipitations that occur here are in the form of
snow. Lakes are characterized by fresh and hard waters (0.2 - 4 mS cm-1; 144 - 1546 mg l-1
CaCO3), of variable pH (between 7.8 and 9.4).

Diatom Assemblages and Their Relationship to Environmental Parameters

Diatom Richness and Diversity
A total of 160 diatom species were identified in the modern assemblages. Among these,
116 taxa (Appendix I) were found in an abundance > 1% in at least one lake and were
retained in numerical analyses (Bigler and Hall, 2002). Richness ranged between 6 (Site 25)
and 41 taxa (Site 3, Figure 7). Shannon diversity values were also highly variable between
sites, ranging between 0.47 (Site 28) and 3.27 (Site 3, Figure 8). The majority of taxa
belonged to Navicula (20 taxa) and Nitzschia (14 taxa) comprising 17.4% and 12.2% of the
total number of taxa, respectively. Cyclotella meneghiniana and Hippodonta hungarica were
the most widely distributed species, being present in 20 and 21 lakes, respectively (Figure 9).

Figure 7. Diatom richness across the study area.

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

Figure 8. Diatom diversity (Shannon H) across the study area.

Detrended Correspondence Analysis (DCA) was performed on diatom assemblages in

order to estimate species turnover (Eilertsen et al. 1990). DCA calculates the length of the
species gradients (DCA axis 1 scores), which describe the degree of species turnover, with
half change in species composition occurring at approximately 1.01.4 standard deviation
(SD) units, and gradient lengths more than 4 SD indicating a complete turnover of species
(Heino and Soininen, 2005). Results for the central Argentina dataset showed strong shifts on
diatom composition along the longitudinal gradient (Figure 9), indicating strong turnovers (up
to 3 S.D.) near the limits of the main ecoregions represented in the dataset (Pampas, Espinal
and Monte/Andean steppe). Aulacoseira granulata, Fragilaria inflata, Navicula veneta,
Cocconeis placentula and Navicula cincta formed an assemblage characteristic of pampean
lakes; while Amphora copulata, Gomphonema parvulum, Pinnularia borealis, Navicula cf.
seminulum and Hantzschia amphyoxis were abundant in lakes from the Espinal region (Figure
9). Nitzschia compressa, Achnanthidium minutissimum, Staurosira pinnata, S. venter, among
others, dominated in lakes located westwards (sites 25 to 34; Figure 9).

Diatom-Environment Ordination
Paleoecological interpretations of changes in the physicochemical characteristics of lakes
can be made qualitatively and/or quantitatively. Although quantitative inferences are always
preferred, there is little point in attempting to quantify a weak environmentspecies
relationship (Smol and Cumming, 2000). Hence, the first step towards the development of a
quantitative reconstruction is to evaluate the strength of the relationship between diatom
assemblage composition and the environmental variables. This can be done through the
application of a number of ordination techniques (ter Braak and Prentice, 1988), from which
Canonical Correspondence Analysis (CCA) is the most commonly applied in diatom research
(ter Braak, 1986). Therefore, the Central Argentina diatom-dataset was related to the

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

195

environmental parameters by CCA using the program CANOCO version 4.5 (ter Braak and
milauer, 1998). Partial CCAs, run with one explanatory variable at each time, were also
applied in order to separate the total variation in diatom data into components that represent
the unique contributions of individual and significant environmental variables (Bocard et al.,
1992).

Figure 9. Distribution of the dominant diatom taxa in surface sediments of the 34 sampling sites.

This approach allowed identifying summer T and pH as the strongest variables explaining
significant portions of the variance within the diatom data set (6.65 and 6.22, respectively;
p<0.001). The other statistically significant environmental variables were SiO2 (5.8%;
p<0.001), gravel (5.7%), hardness (5.1%), log-conductivity (4.8%), vegetation cover (4.7%),
and coarse sand (4%; p<0.05). This reduced set of environmental variables explained 35.6%
of the variation in diatom distribution and produced highly significant canonical axes
(p<0.001). The species-environment correlations of CCA axis 1 (0.953) and axis 2 (0.876)
were high and indicated a strong relationship of diatom relative frequencies to environmental
variables (Table 2).

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

Table 2. Summary statistics for the first two axes of CCA, with the nine selected
environmental factors.
Axis

Ordination results
1
2
0.500
0.399
0.953
0.876

Eigenvalues ()
Species-environment correlation
Cumulative % variation of:
Species data
7.6
Species-environment relationship
21.6
Sum of all
6.552
Sum of all canonical
2.316
Test of significance of the first canonical axis p<0.005
Test of significance of all canonical axis p<0.001

13.7
38.8

The CCA ordination plot showed that composition of surface sediment assemblages
differed between the main ecoregions (Figure 10). The first axis (7.6% of the explained
variance) showed a positive correlation with SiO2 (r = 0.72), gravel (r = 0.57) and vegetation
cover (r = 0.54) and a negative correlation with summer T (r = -0.82) and pH (r = -0.73).
Accordingly, sites were ordinated along this axis following mainly temperature, pH and
silicates gradients, with shallow lakes from the Espinal and Monte located towards the left
and right sides of the plot, respectively. The second axis (6.1% of the variance explained) was
positively correlated with water hardness (r = 0.67), and log-conductivity (r = 0.60).
Consequently, sites were ordinated along axis 2 following ionic strength gradients, with
saline and brackish lakes located towards the top section of the diagram (Figure 10).

Figure 10. CCA ordination plot of diatom assemblages against selected environmental variables.

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

197

Diatom-Based Transfer Functions

In the last decades, paleoecologists have cataloged the ecological optima and tolerances
of diatom assemblages and have constructed transfer functions that allow environmental
conditions to be inferred from fossil assemblage data (Birks et al., 1990). Transfer functions
simply describe the relationship between diatom assemblages and their environment with the
ultimate goal to use the present-day ecology of diatoms to quantitatively infer past conditions.
This approach involves two steps: (1) the regression step in which the responses of taxa to the
contemporary environment are modeled, and (2) a calibration step in which the
environmental variable of interest is predicted from the fossil compositional data (Birks,
1995). The mathematical methods for establishing a transfer function are commonly based on
nonlinear regression techniques. The most popular approaches in diatom studies have been
Weighted Averaging (WA; ter Braak and van Dam, 1989) and the WA partial least squares
(WAPLS; ter Braak and Juggins, 1993) regression, because they perform well with noisy,
species-rich, compositional data, with many zero values and some outliers. In both methods,
the underlying response model is the Gaussian function, that is, each taxon has a bell shape
distribution along an environmental variable. From the Gaussian curve, the species optimum
(the peak) and its tolerance (the standard deviation) can be estimated (Korhola, 2007).
Results of CCA ordination suggests that diatoms from Central Argentina are potential
indicators of environmental variables directly (summer T) and indirectly (pH and logconductivity) related to climate. After detecting these strong and significant relationships
between diatom assemblages and environmental variables, it is possible to model this
relationship through transfer functions. Hence, various calibration models were tested for the
training set using the program C2 1.6.0 (Juggins, 2003). The performance of each model was
addressed by analyzing the different statistical parameters produced by C2: coefficient of
determination (r2), apparent root mean square error (RMSE), mean and maximum bias. As
RMSE is under-estimated when based solely on the training set, and because no independent
test set was available for this study, a cross-validation procedure known as jack-knifing was
applied, in order to derive a more realistic and reliable estimate of the overall RMSE of
prediction for the training set (RMSEP jack-knifing). Ideally, the best models should have high r2
and low RMSEP when compared to the variable range, and low mean and maximum bias
based on cross-validation (ter Braak and Juggins, 1993). Diatom taxon responses (optima and
tolerances) were modeled along the salinity gradient using WA with C2 1.6.0 (Juggins, 2003).
Weighted-averaging salinity optima and tolerances of diatom taxa were calculated for
pH, summer temperature and log-conductivity (Figures 11 to 13, Appendix I), as well as the
number of occurrences of each taxon (N) and Hill's N2 (number of effective occurrences; Hill,
1973). The latter gives an indication of the number of samples contributing to the calculation
of a taxon's WA optima. Optima estimated for taxa with a low number of occurrences or a
low N2 should be interpreted with caution (Gasse et al., 1995). Unfortunately, a large number
of taxa in the training set showed low N2 (Appendix I). According to the performance of the
several transfer functions developed (Table 3), the two-component WAPLS models appear
to perform better than WA for the three variables modeled when examining the apparent r2
(pH=0.96, summer temperature=0.97, log-conductivity=0.93), RMSE (pH=0.17, summer
temperature=0.58, log-conductivity=0.11), and maximum deviation (pH=0.17, summer
temperature=0.70, log-conductivity=0.26; Table 3). Residuals for pH, summer T and logconductivity were within 0.41, 1.6 and 0.26 of the observed values, respectively (Figure
14). However, results of jack-knifing cross-validation revealed significant drops in the

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

performance of these models (Table 3), which showed low r2jack-knifed (pH=0.42, summer
temperature=0.51, log-conductivity=0.20), higher RMSEPjack-knifed (pH=0.65, summer
temperature=2.38, log-conductivity=0.39), and higher maximum deviation (pH=0.85,
summer temperature=2.72, log-conductivity=1.10) based on cross-validation.
Overall, our transfer functions showed low performances under cross-validation when
compared to models developed in other parts of the world. In the case of pH, the reported
r2jack-knifed or boot usually ranged between 0.19 and 0.61, and is comparable to the obtained in the
Central Argentina dataset (0.42). The RMSEP jack-knifed or boot represented the 22.17% of the pH
gradient, being higher than the reported for previous pH transfer functions (7.3-14.4%; Gasse
et al., 1995; Weckstrm et al., 1997; Rosen et al., 2000; Reavie and Smol, 2001; Bigler and
Hall, 2002; Philibert and Prairie, 2002; Denys, 2006; Ginn et al., 2007). For other summer air
temperature transfer functions, the r2jack-knifed or boot ranged between 0.36 and 0.79, which is
comparable with our value (0.51), while the RMSEP jack-knifed or boot represented the 10.7-18.2
% of the total gradient (Pienitz et al., 1995; Lotter et al., 1997; Korhola et al., 2000; Rosn et
al., 2000; Bigler and Hall, 2002), while in our work it represented the 20.32%. Finally, our
conductivity transfer function showed a low RMSEP jack-knifed (14% of the total gradient), but
its performance was very low in terms of r2jack-knifed (0.2). In fact, reported r2jack-knifed values
range between 0.52 and 0.92 (e.g. Gasse et al., 1995; Reed, 1998; Gregory-Eaves et al., 1999;
Davies et al., 2002; Ryves et al., 2002; Yang et al., 2003).
Table 3. Main statistics of the several salinity transfer functions developed for the
calibration set.

r
A) pH
WAINV
WACLA
WATINV
WATCLA
WAPLS-1
WAPLS-2
B) Summer Temperature
WAINV
WACLA
WATINV
WATCLA
WAPLS-1
WAPLS-2
C) log-Conductivity
WAINV
WACLA
WATINV
WATCLA
WAPLS-1
WAPLS-2

Apparent
Max.
RMSE
Dev.

Jack-Knifed
2

Max.
Dev.

RMSEP

RMSEP as
% of gradient

0.82
0.82
0.85
0.85
0.82
0.96

0.32
0.25
0.40
0.48
0.32
0.17

0.36
0.39
0.32
0.35
0.36
0.17

0.49
0.51
0.46
0.46
0.49
0.42

0.79
0.64
0.72
0.60
0.80
0.85

0.60
0.62
0.64
0.68
0.60
0.65

20.39
21.02
21.75
23.10
20.33
22.17

0.87
0.87
0.92
0.92
0.87
0.97

1.20
0.82
1.65
1.29
1.19
0.70

1.21
1.29
0.94
0.98
1.20
0.58

0.49
0.51
0.41
0.42
0.51
0.51

3.05
2.32
4.25
3.89
3.15
2.72

2.39
2.46
2.60
2.64
2.32
2.38

20.51
21.03
22.22
22.53
19.89
20.32

0.80
0.80
0.82
0.82
0.80
0.93

0.48
0.45
0.48
0.50
0.49
0.26

0.19
0.21
0.18
0.20
0.19
0.11

0.28
0.31
0.17
0.19
0.29
0.20

1.03
0.90
1.03
0.90
1.12
1.10

0.37
0.37
0.40
0.41
0.36
0.39

13.73
13.89
14.99
15.47
13.50
14.58

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

Figure 11. pH optima and tolerances for the dominant diatom taxa in the training set.

199

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

Figure 12. Maximum summer temperature optima and tolerances for the dominant diatom taxa in the
training set.

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

201

Figure 13. Conductivity (log10 transformed) optima and tolerances for the dominant diatom taxa in the
training set.

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

Figure 14. Plots of observed versus inferred values and observed versus residual values based on WAPLS regression and calibration for pH (top), summer T (middle) and log-conductivity (bottom).

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

203

The poor performances under jackknifing are usually a result of the heterogeneous nature
of the data and the relatively small size of the dataset, such that there are many samples
dominated by rare taxa (which is in agreement by the low number of occurrences showed by
many taxa in our dataset, Appendix I), making these samples poorly predicted when left out
of the training set (Reed, 1998). Moreover, the environmental gradients, particularly the
conductivity one, are not fully covered in our dataset (Figure 14). This constitutes a problem,
since it is well known that transfer functions are sensitive to the distribution of the
environmental variables in the training set, so that the method performs well only when
samples have a reasonably even distribution along the chemical gradient and when species are
sampled over their whole range (ter Braak, 1986). These limitations highlight the benefits of
using large datasets, which provide a more thoroughly representation of the environmental
gradients and, as such, of the ecology of species assemblages (Ginn et al., 2007). Hence,
further improvement in model performance is expected as more lakes are added to our initial
data-set.

PALEOCLIMATOLOGICAL SIGNIFICANCE
The biggest difficulty when applying transfer functions to the fossil record is finding
training sets representative of the fossil assemblages. Besides the requirement of a statistical
robust model, a good agreement between modern and fossil diatom assemblages is also a
major requisite in order to apply transfer functions (Birks, 1995). The lack of analogy
between modern and fossil diatom assemblages prevents the transfer of modern
environmental information to the past, restricting the applicability of quantitative models.
Thus, a reliable reconstruction must be based on good modern analogues (Hbener et al.,
2008).

Figure 15. Distribution of the dominant diatom taxa along the Nahuel Ruc core (modified after Stutz et
al., in press).

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G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

In order to assess the potential use of our modern data set as analogues of past diatom
assemblages in pampean shallow lakes, fossil samples were compared to modern samples
from a core obtained with a livingston corer in the deepest part of Nahuel Ruc lake (Stutz et
al., in press). The core is 123 cm long, and covered the last ca. 3700 cal years BP (Figure 15).
Seventy-four diatom species were identified throughout the sediment core, and no valves
were preserved below 82 cm. According to diatom assemblage composition, four diatom
zones were defined (Stutz et al., in press). From 82 to 69 cm (Zone I), assemblages were
dominated by Aulacoseira granulata var. angustissima, Campylodiscus clypeus, A. granulata,
and Cyclotella meneghiniana. The zone II, between 69 and 55 cm, was dominated by C.
meneghiniana and E. adnata, together with Aulacoseira muzzanensis, A. granulata and
Fragilaria capuccina. Between 55 and 21 cm (Zone III) a sharp increase in the relative
abundance of A. granulata, accompanied by high abundances of C. meneghiniana and
Aulacoseira muzzanensis was registered. The top of the sequence (200 cm) exhibited the
highest relative abundances of E. adnata, Cocconeis placentula, and Rhopalodia gibberula,
together with A. granulata and C. meneghiniana, and peaks of A. granulata var. angustissima
and Pleurosira laevis.

Figure 16. Results of analog analysis (MinDC), diatom inferred curves of pH and summer T and
phytolith temperature index (modified after Stutz et al., in press) based on Nahuel Ruc core.

The comparison between modern and fossil diatom assemblages was based in the Modern
Analogue Technique (MAT), a technique useful to detect fossil samples that have close

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

205

modern analogues within the available modern training set. The basic idea is to compare
numerically, using the Squared Chord Distance (SCD) as the dissimilarity measure, the
diatom assemblage in fossil assemblages with the ten most similar modern samples. Fossil
samples with coefficients below the tenth percentile of the modern training set are considered
to have good analogues (Birks et al., 1990). Transfer functions developed for pH and summer
temperature were applied to the core in order to reconstruct past changes in these two
environmental variables, and the results were compared to the qualitative environmental
inferences based on a multi proxy approach provided by Stutz et al. (in press). Both MAT and
transfer function analyses were developed using C2 version 1.4.3 (Juggins, 2003).

MAT
In general, a good agreement between modern and fossil diatom assemblages was found.
Examination of the MAT results showed that the SCD values were generally below the tenth
percentile of the training set samples (Figure 16), indicating a good match between fossil and
modern samples. Three samples from zone II and five from zone IV remained without a
modern equivalent (Figure 16). All these samples contained high relative frequencies of
Epithemia adnata, which was rarely found in the modern training set, appearing in low
frequencies (<2%) in some sites (Los Pinos and El Sosneado lakes). However, WA appears to
perform well in "no-analogue" situations where the fossil assemblages differ, to some degree,
in composition to the modem assemblages, since the WA inferences are based on the
weighted average of the individual optima of the taxa in common between the modern and
fossil assemblages (Birks and Birks, 1980; Birks, 1995). Hence, the transfer functions can be
applied to these samples, but the inferred values must be interpreted with caution.

Transfer Function Application and Comparison with Other Indicators

The application of the jack-knifed two-component WAPLS models previously
developed allowed the quantitative reconstruction of pH and summer temperature based on
fossil diatom assemblages from Nahuel Ruc core. According to the inferred pH curve, pH
values remained relatively constant all along the sequence, ranging between 8.2 and 8.9
(Figure 16). The inferred curve of summer temperature, on the other hand, showed significant
fluctuations along the core (Figure 16). Inferred values for the base were high (312.4C) and
showed a decreasing tendency until reaching its lowest value (25.22.4C) at a depth of 68
cm. Values increased again along zone II, reaching a value of 292.4C at a depth of 50 cm.
The inferred temperature showed a decreasing tendency along the rest of the sequence,
reaching 25.52.4C at a depth of 13 cm. In the last 13 cm, however, this tendency reversed,
and temperature increased again reaching 302.4C at the top of the core. This last value fits
well with the maximum values of January temperature reported by instrumental data for the
near Mar del Plata city (e.g. 33.4C for January 2010).
The curve of reconstructed changes in summer temperature agreed quite well with the
plotted temperature index based on phytolith assemblage composition (Figure 16). This index
is based in the ratio of chloridoid phytoliths to the sum of chloridoid and panicoid phytoliths,
and allows and indirect inference of changes in mean annual temperature. High index values
indicate an arid climate, while low values indicate a humid climate. Opal phytoliths in surface

206

G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

deposits of cattle/sheep enclosures registered Humidity Index (HI) values between 10 in

southern Uruguay and 16 (SE Buenos Aires Province) for Pampa grasslands (del Puerto,
2009).
Based on this index, Stutz et al. (in press) suggested warm and humid conditions at the
beginning of the late Holocene, after ca 3,700 cal year BP, with an episode of cooler and
dryer conditions at ca. 740 cal years BP. During the last 400 cal years, the phytolith
temperature index indicates a decrease in temperature, which was associated with the Little
Ice Age. The main differences between diatom and phytolith inferred temperature curves are
recorded at the last 10 cm of the core, where phytoliths indicate a moment of higher
temperature not represented in the diatom curve. Moreover, according to this index, cold and
humid-dry conditions persisted at Nahuel Ruc until present, with no register of the
increasing temperature values suggested by the diatom transfer function for the top 5 cm of
the core (Figure 16). Addressing if this last temperature increase is an artifact or a real
temperature change requires further study.

DISCUSSION AND CONCLUSIONS

The significant relationships found between diatoms and climatically-driven
environmental variables encourage the application of diatom-based quantitative methods for
reconstructing the Quaternary climate of Central Argentina. The low to very low
performances obtained for the preliminary developed transfer functions can be attributed to
the low number and the heterogeneous nature of the sites in our data set. The problems
associated with relatively small, regional data-sets, such as uneven distribution of sites along
the environmental gradients of interest and truncated species distributions (see e.g. Bennion et
al., 1996; Weckstrm et al., 1997), are evidently present in our study. A future expansion of
the training-set to at least 100 shallow lakes will probably extend the pH, temperature and
conductivity gradients, increase the number of occurrences of dominant taxa, and improve the
performance of the calibration models, which are known benefits of large datasets (Ginn et
al., 2007).
Summer temperature was the strongest environmental variable explaining a significant
portion of the variance of diatom assemblages in our dataset, which encouraged the
development of a diatom based air temperature transfer function. However, the relationship
between diatom distribution and temperature is still poorly understood, and consequently
diatom-based temperature transfer functions are considered with uncertainty (Anderson,
2000). In addition, there are many potentially strong physical (habitat), chemical (pH,
nutrients) and biological (food web effects) influences on diatom assemblages that may act
independently of temperature (Battarbee, 2000). Despite the doubt about diatom-temperature
transfer function performance, several such models have been developed from training sets in
several regions of the world (Pienitz et al., 1995; Weckstrm et al., 1997; Rosn et al., 2000;
Bigler et al., 2002; Vyverman and Sabbe, 1995; Lotter et al., 1997). In fact, this development
has been encouraged because performance statistics of the diatom-temperature transfer
functions are usually comparable to those based on other proxy-indicators (e.g., chironomids,
pollen) from the same training-sets (e.g., Lotter et al., 1997; Rosn et al., 2000). However,
the relationship between diatoms and climate may be complex and probably temperature is

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

207

related to a combination of several factors (Pienitz et al., 1995; Lotter et al., 1997). As a
consequence, the diatoms estimated optima and tolerances for air temperature should not be
considered as absolute values, but also as relative to the other taxa as well as other indicators.
The observed strong relationship between pH and diatoms is in agreement with results of
numerous previous studies (e.g. Gasse and Tekaia, 1983; Davis and Anderson, 1985; Dixit et
al., 1988; Weckstrm et al., 1997; Bigler and Hall, 2002; Denys, 2006; Ginn et al., 2007). As
diatoms respond sensitively to changes in pH, diatom pH transfer functions have been
commonly applied in acidification studies (e.g. Davis et al., 1983; Battarbee and Charles,
1987; Wckstrom et al., 2003). More recently, this approach has also been used in climate
change reconstruction (Boyle, 2008).
Diatom-inferred salinity/conductivity methods are now being used extensively, separately
or in combination with other proxy methods, for climate change studies (Battarbee, 2000).
Diatoms are especially good indicators of salinity, and have been used extensively for the
purposes of climate reconstruction from closed basin lake sediments (Bradbury, 1989;
Bradbury et al., 1981; Gasse, 1987; Gasse et al., 1997). As the ionic concentration and
composition of inland water bodies are affected by changes in effective moisture
(precipitation minus evaporation: PE), changes in the composition of diatom species that
vary in their salinity tolerances provide a tool for the reconstruction of past climate (Fritz,
2007). Hence, in the last decade several diatom-salinity training sets for different
geographical regions have been constructed for paleoclimatological purposes (Fritz et al.,
1991; Gasse et al., 1995; Wilson et al., 1996; Gell, 1997; Reed, 1998). In the case of our
training set, the conductivity model showed a very low performance and could not be applied
to the fossil record. Hence, further study is needed in order to increase its predictive
capability and allow diatom-based conductivity reconstructions in Central Argentina.
Overall, the strong statistical relationship between diatoms and air temperature, pH and
conductivity suggest the potential application of diatom based models to the quantitative
reconstruction of these variables during the Holocene in Central Argentina. Moreover, our
reduced training set showed to be good analogs of fossil assemblages from Nahuel Ruc lake,
increasing the good perspectives for quantitative reconstruction. However, more work is
needed in order to produce significant and reliable reconstructions; the addition of new lakes
to the database, covering the full range of variability of the environmental variables selected,
is a critical next step to do.

REVIEWED BY
Dr. Marcela Tonello, Universidad Nacional de Mar del Plata, Departamento de
Biologa, CP: 7600, Mar del Plata, ARGENTINA, e-mail: mtonello@mdp.edu.ar.
Dr. Silvina M. Stutz, Universidad Nacional de Mar del Plata, Departamento de Biologa,
CP: 7600, Mar del Plata, ARGENTINA, e-mail: silstutz@yahoo.com.ar.
Lic. Alejandra Escandell, Universidad Nacional de Buenos Aires, Ciudad Universitaria.
Pabelln II. CP: C1428EHA. Buenos Aires, ARGENTINA, e-mail: alescan21@hotmail.com.

208

G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

ACKNOWLEDGMENTS
We are very grateful to Frank Columbus for inviting us to participate of this publication.
We thank Felipe Garca-Rodriguez and Silvina Stutz for gently providing the diatom data set
from the Nahuel Ruc core. Marcela Tonello, Silvina Stutz and Alejandra Escandell made
helpful commentaries and suggestions on the manuscript. Financial support for this work was
provided by the Conservation Research and Education Opportunities (CREO), University of
Mar del Plata (EXA 457/09), and CONICET (PIP 0063/2010). G.S.H. and C.G.D.F. are
members of the Scientific Research Career of the Consejo Nacional de Investigaciones
Cientficas y Tcnicas (CONICET). E.T. and P.A.C. were founded by CONICET fellowships.

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215

APPENDIX I
Number of occurrences (N) and effective number of occurrences (Hill's N2) of the diatom
taxa, their Weighted-Averaging optima (Opt.) and tolerances (Tol.) for summer T, pH and
log-Conductivity. Numbers correspond to the ones used in CCA.
Taxon Name
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50

Achnanthes cf. manguinii

Achnanthes exigua
Achnanthidium minutissimum
Adlafia aquaeductae
Amphora coffeaeformis
Amphora cognata
Amphora copulata
Amphora delicatissima
Amphora pediculus
Amphora sp.
Amphora veneta
Anomoeoneis sphaerophora
Aulacoseira granulata
A granulata var. angustissima
Caloneis westii
Cocconeis placentula
Coscinodiscus sp.
Ctenophora pulchella
Cyclotella meneghiniana
Cyclotella ocellata
Cymbella affinis
Denticula elegans
Diatoma moniliformis
Diatoma vulgare
Dickieia subinflata
Encyonema minutum
Encyonopsis microcephala
Eolimna minima
Epithemia adnata
Epithemia sorex
Eunotia pectinalis
Fallacia pygmaea
Fragilaria capucina
Fragilaria fasciculata
Fragilaria inflata
Fragilaria sp.
Fragilariforma virescens
Gomphoneis herculeana
Gomphonema clavatum
Gomphonema gracile
Gomphonema parvulum
Gomphonema truncatum
Hantzschia amphyoxis
Hantzschia virgata
Hippodonta hungarica
Karayevia clevei
Lemnicola hungarica
Luticola cohnii
Luticola kotschyi
Luticola mutica

Taxon Authority
Hustedt
Grunow
(Ktzing) Czarnecki
(Krass)Mos.LBert&Met
(Agardh) Ktzing
Cholnoky
(Ktz.) Schoe & Arch
Krasske
(Ktzing) Grunow
Ktzing
Pfitzer
(Ehrenberg) Simonsen
(Mller) Simonsen
(Smith) Hendey
Ehrenberg
(Ralfs) Will & Round
Ktzing
Pantocsek
Ktzing
Ktzing
Ktzing
Bory de Saint-Vincent
(Grunow) D.G.Mann
(Hilse) D.G.Mann
(Grunow) Krammer
(Grun.) Lange-Bertalot
(Ktzing) Brbisson
Ktzing
(Ktzing) Rabenhorst
(Ktz) Stickle & Mann
Desmazires
(Agardh) L-Bertalot
(Heiden) Hustedt
(Ralfs) Will. & Round
(Ehrenberg) Cleve
Ehrenberg
Ehrenberg
(Ktzing) Ktzing
Ehrenberg
(Ehrenberg) Grunow
(Roper) Grunow
(Grun)Bert.Metz.&Wit.
(Grun) Round & Bukh
(Grun) Round & Bass.
(Hilse) Mann
(Grunow) Mann
(Ktzing) Mann

N
1
3
14
1
15
2
13
1
6
1
18
10
7
6
4
19
1
6
20
2
3
5
4
3
1
4
2
5
6
7
1
2
7
1
2
1
4
1
7
3
11
5
12
1
21
1
2
2
2
6

N2
1.00
2.44
7.90
1.00
12.25
1.80
7.01
1.00
3.85
1.00
13.8
7.05
5.96
4.72
2.55
12.31
1.00
4.04
13.71
2.00
2.72
4.30
1.98
2.44
1.00
3.88
2.00
4.65
4.98
5.18
1.00
1.90
5.08
1.00
1.93
1.00
3.23
1.00
4.15
2.83
6.72
2.83
6.73
1.00
14.91
1.00
1.37
1.49
1.74
4.93

Summer T
Opt
Tol
32.7
2.42
30.97
2.14
28.02
4.61
27.10
2.42
29.96
2.87
27.63
1.63
31.55
1.88
33.60
2.42
27.99
3.99
29.60
2.42
29.15
3.05
30.44
2.24
29.49
1.54
29.87
1.63
31.70
1.82
28.47
2.70
30.60
2.42
27.32
3.60
30.65
2.24
29.18
1.20
28.00
3.36
28.38
1.83
27.48
3.20
24.80
4.47
32.00
2.42
25.12
3.45
29.19
1.20
27.34
4.51
25.93
3.33
25.7
4.45
27.1
2.42
27.06
0.07
25.95
3.24
26.1
2.42
30.01
1.56
23.7
2.42
26.38
3.26
22.00
2.42
27.27
2.54
31.46
2.09
30.42
3.61
30.13
4.57
32.27
1.52
28.40
2.42
30.52
2.54
22.00
2.42
27.13
0.14
32.16
0.14
27.24
0.14
28.99
1.86

pH
Opt
10.5
9.77
8.66
8.60
9.16
7.93
9.85
8.26
8.11
9.02
9.12
9.60
8.96
8.76
9.44
8.70
9.02
8.41
9.46
7.87
7.83
8.59
7.94
7.92
9.20
8.76
7.87
9.15
8.55
8.99
8.60
8.75
8.50
8.04
8.93
7.81
8.45
9.00
8.22
9.48
9.47
9.68
9.80
8.25
9.38
9.00
8.68
9.68
8.96
8.72

Tol
0.59
0.52
1.00
0.59
1.02
0.12
0.71
0.59
0.65
0.59
0.84
0.75
0.32
0.38
0.96
0.83
0.59
0.67
0.68
0.23
0.20
0.63
0.20
0.61
0.59
0.51
0.23
0.97
0.74
0.43
0.59
0.27
0.57
0.59
0.44
0.59
0.63
0.59
0.44
1.05
0.88
1.05
0.46
0.59
0.68
0.59
0.37
0.42
0.37
0.66

log-Cond
Opt
Tol
0.45
0.29
0.82
0.34
0.38
0.30
0.33
0.29
0.76
0.36
0.89
0.20
0.61
0.29
1.03
0.29
0.27
0.11
0.74
0.29
0.55
0.29
0.67
0.30
0.53
0.25
0.40
0.14
0.59
0.36
0.53
0.33
0.42
0.29
0.44
0.42
0.54
0.27
0.28
0.07
0.27
0.06
0.55
0.28
0.31
0.06
0.22
0.07
2.00
0.29
0.21
0.14
0.28
0.07
0.28
0.16
0.40
0.34
0.29
0.28
0.33
0.29
0.53
0.36
0.25
0.12
0.70
0.29
0.47
0.45
0.25
0.29
0.57
0.47
0.09
0.29
0.32
0.16
0.58
0.36
0.52
0.33
0.66
0.43
0.52
0.32
0.31
0.29
0.58
0.35
0.09
0.29
0.32
0.08
1.14
0.77
0.25
0.08
0.46
0.24

G. S. Hassan, E. Tietze, C. G. De Francesco, and P.A. Cristini

216
Taxon Name
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109

Luticola sp.
Mastogloia braunii
Mastogloia elliptica
Mastogloia pumila
Melosira varians
Navicella pusilla
Navicula capitellata
Navicula cari
Navicula cf. angusta
Navicula cf. rhynchocephala
Navicula cf. seminulum
Navicula cincta
Navicula cryptocephala
Navicula erifuga
Navicula gregaria
Navicula kefvingensis
Navicula libonensis
Navicula microdigitoradiata
Navicula peregrina
Navicula perminuta
Navicula recens
Navicula rostellata
Navicula sp. 1
Navicula sp. 2
Navicula tripunctata
Navicula veneta
Nitzschia amphibia
Nitzschia austriaca
Nitzschia cf. palea
Nitzschia communis
Nitzschia compressa
Nitzschia filiformis
Nitzschia fonticola
Nitzschia inconspicua
Nitzschia linearis
Nitzschia microcephala
Nitzschia perminutum
Nitzschia scalpelliformis
Nitzschia sp.
Nitzschia umbonata
Opephora sp.
Pinnularia borealis
Pinnularia sp.
Pinnularia subcapitata
Pinnularia viridis
Planothidium delicatulum
Planothidium lanceolatum
Pleurosira laevis
Pseudostaurosira brevistriata
Reimeria sinuata
Rhoicosphenia abbreviata
Rhopalodia constricta
Rhopalodia gibba
Rhopalodia gibberula
Rhopalodia musculus
Stauroneis tackei
Staurosira construens
S. construens var. subsalina
Staurosira venter

Taxon Authority
Grunow
(Agardh) Cleve
(Cleve) Cleve
Agardh
(Grunow) Krammer
Cleve
Ehrenberg
Grunow
Ktzing
Grunow
(Ehrenberg) Ktzing
Ktzing
Lange-Bertalot
Donkin
(Ehrenberg) Ktzing
Schoeman
Lange-Bertalot
(Ehrenberg) Ktzing
Grunow
(L-Bert) L-Bertalot
Ktzing
(Mller) Bory
Ktzing
Grunow
Hustedt
(Ktzing) W.Smith
Rabenhorst
(Bailey) Boyer
(Smith) Hustedt
(Grunow) Grunow
Grunow
(Agardh) Smith
Grunow
(Grunow) Peragallo
Grunow
(Ehr) Lange-Bertalot
Ehrenberg
Gregory
(Nitzsch) Ehrenberg
(Ktz) Round & Bukh
(Brb) Lange-Bertalot
(Ehrenberg) Compre
(Grun) Will & Round
(Greg) Koc& Stoermer
(Agardh) Le-Bertalot
(Smith) Krammer
(Ehrenberg) Mller
(Ehrenberg) Mller
(Ktzing) Mller
(Hust) Kram & L-Bert.
Ehrenberg
(Hust) Haw. & Kelly
(Ehr) Kobayasi

N
3
1
6
1
3
6
1
4
1
2
8
13
2
5
5
3
4
1
12
3
4
2
6
1
2
13
14
8
5
3
5
1
2
12
2
9
7
1
1
3
1
13
2
2
2
8
4
2
11
1
7
1
4
7
2
1
1
1
13

N2
2.56
1.00
4.58
1.00
2.73
4.48
1.00
2.53
1.00
1.65
5.28
6.10
1.45
2.84
4.04
2.41
3.52
1.00
9.09
2.91
1.76
1.89
4.49
1.00
1.95
8.47
8.32
7.00
4.64
2.31
2.84
1.00
1.76
10.08
1.80
5.72
4.98
1.00
1.00
2.30
1.00
8.83
1.83
1.73
1.88
4.25
3.57
1.78
6.61
1.00
4.46
1.00
3.12
5.09
1.57
1.00
1.00
1.00
5.66

Summer T
Opt
Tol
32.52
0.26
28.40
2.42
27.39
1.15
28.40
2.42
26.28
3.24
27.31
1.25
32.20
2.42
32.53
0.52
32.40
2.42
27.08
0.21
32.69
0.57
28.87
2.86
32.65
0.92
29.74
1.43
28.66
4.12
27.12
0.13
25.58
2.66
32.00
2.42
30.58
1.98
27.49
0.79
29.59
5.57
26.55
0.85
30.77
3.00
28.30
2.42
28.68
2.33
28.05
2.45
30.27
3.08
27.98
2.98
27.65
0.62
31.00
4.25
30.51
3.05
31.6
2.42
22.22
0.49
27.91
3.32
25.21
6.79
26.79
2.92
25.71
3.82
27.40
2.42
27.30
2.42
32.52
0.23
28.40
2.42
32.38
1.32
32.89
0.64
26.46
4.53
27.87
0.99
30.04
3.20
24.39
3.24
31.37
0.49
26.64
3.53
30.00
2.42
28.03
3.77
32.00
2.42
32.43
0.77
26.45
2.66
30.81
3.54
27.3
2.42
28.70
2.42
28.70
2.42
26.13
3.99

pH
Opt
10.2
7.87
8.53
7.87
7.90
8.35
9.80
9.84
9.70
9.02
9.76
8.69
9.78
8.88
9.07
8.80
8.63
9.20
9.18
8.64
9.00
8.45
9.64
9.80
9.08
8.95
9.15
8.65
9.06
8.13
8.13
9.04
8.60
8.76
9.28
8.52
8.60
8.25
9.12
10.09
7.87
9.68
10.15
8.60
8.52
9.31
8.50
8.92
8.82
7.71
8.45
9.20
9.87
8.44
9.15
9.12
9.30
9.30
8.26

Tol
0.44
0.59
0.52
0.59
0.24
0.56
0.59
0.35
0.59
0.10
0.36
0.72
0.28
0.77
0.79
0.30
0.89
0.59
0.65
0.39
0.61
0.76
0.55
0.59
0.07
0.70
0.93
0.77
0.57
0.21
0.88
0.59
0.91
0.62
0.25
0.72
0.90
0.59
0.59
0.51
0.59
0.66
0.71
0.81
0.52
0.80
0.58
0.25
0.61
0.59
0.76
0.59
0.44
0.56
0.16
0.59
0.59
0.59
0.98

log-Cond
Opt
Tol
0.38
0.25
0.99
0.29
0.68
0.25
0.99
0.29
0.60
0.45
0.75
0.24
0.91
0.29
0.35
0.46
0.08
0.29
0.67
0.44
0.48
0.35
1.17
0.70
0.2
0.43
0.43
0.23
0.31
0.30
0.42
0.30
0.39
0.30
2.00
0.29
0.44
0.29
0.53
0.31
0.42
0.26
0.52
0.34
0.29
0.21
0.84
0.29
0.30
0.15
0.56
0.27
0.49
0.27
0.79
0.63
0.56
0.32
0.96
0.18
1.21
0.35
0.45
0.29
0.14
0.11
0.42
0.29
0.21
0.26
0.57
0.3
0.40
0.35
0.31
0.29
0.22
0.29
0.29
0.27
0.99
0.29
0.55
0.45
0.31
0.06
0.24
0.16
0.51
0.37
0.68
0.55
0.21
0.13
0.37
0.17
0.54
0.43
0.23
0.29
0.25
0.13
2.00
0.29
0.49
0.28
0.55
0.28
1.72
0.82
0.22
0.29
0.85
0.29
0.85
0.29
0.41
0.34

Problems and Potentialities of Using Diatoms as Paleoclimatic Indicators

Taxon Name
110
111
112
113
114
115
116

Staurosirella pinnata
Stephanodiscus niagarae
Surirella minuta
Surirella ovalis var. linearis
Surirella striatula
Synedra acus
Tabularia tabulata

Taxon Authority
(Ehr) Will. & Round
Ehrenberg
Brbisson
Peragallo
Turpin
Ktzing
(Agardh) Snoeijs

N
9
2
2
2
5
7
4

N2
4.79
1.22
1.46
1.77
4.11
5.63
2.63

Summer T
Opt
Tol
24.56
3.54
30.70
0.71
32.06
4.10
28.80
5.30
29.09
2.15
24.32
2.82
26.19
2.37

pH
Opt
8.84
9.02
10.06
9.07
9.26
8.50
8.22

Tol
0.76
0.01
1.59
1.31
0.48
0.64
0.48

217

log-Cond
Opt
Tol
0.37
0.44
0.43
0.02
0.29
0.02
0.71
0.47
0.92
0.54
0.24
0.19
0.38
0.25