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Wetlands: Monitoring, Modelling and Management – Okruszko et al. (eds)
© 2007 Taylor & Francis Group, London, ISBN 978-0-415-40820-2
Testate amoebae as ecological and palaeohydrological indicators
in peatlands – The Polish experience
Mariusz Lamentowicz
Department of Biogeography and Palaeoecology, Adam Mickiewicz University, Pozna´n, Poland
EdwardA.D. Mitchell
Ecole Polytechnique Fédérale de Lausanne (EPFL), Laboratory of Ecological Systems, Lausanne, Switzerland
Institut Fédéral de Recherches WSL, Antenne Romande, Lausanne, Switzerland
ABSTRACT: In this paper we present brief overviews of the ecological study of peatland testate amoebae in
northern Poland and applications of testate amoeba analysis in palaeohydrology as well as for the monitoring of
naturaland human-impacted mires –e.g. Polish Baltic bogs.Testateamoebae are unicellularorganisms that pro-
duce a test (shell), which protects their cytoplasm.These protists represent a valuable tool in palaeohydrological
studies in Sphagnum mires. In peatlands, testate amoebae live in mosses and the upper, oxygenated part of the
peat. They are very sensitive to water table changes and, to a lesser extend also to pH. Some species occur in
moist, slightly acidic hollows and others in dry very acidic Sphagnum hummocks. During the vegetation season
testate amoebae produce many generations. When conditions become less favourable, (e.g. winter or temporary
desiccation in summer), they encyst or die. Empty tests are preserved in peat together with the remains of plants
and some other organisms and become part of the peat archive. Peat sediments provide a unique opportunity
to reconstruct past hydrological changes in mires on the basis of testate amoebae. To reconstruct water table
changes quantitatively a good modern data set is required. For Poland such a data set was recently created from
surface samples from Tuchola Pine Forest. We modelled the response of species (optimum and tolerance) to
environmental variables. Subsequently, this training set was used for inference of past hydrological conditions
from Sphagnum mires in northern Poland,Tuchola and JeleniaWyspa located in theTuchola sandr area. In addi-
tion, plant macrofossils and palynological analyses were used to reconstruct changes in the local and regional
vegetation and the history of human impact. In both sites we observed correlations between the testate amoebae
inferred hydrology and climate changes as well as human activities (e.g. deforestations or damming).
1 INTRODUCTION
Testate amoebae (Protista, also referred to as Tes-
tacea, testaceans, Arcellaceans,) are characterised by
the presence of a shell (called test) that protects the
cell and that allows identification. Over 70 taxa have
so far been found in mires of Poland (Lamentowicz &
Mitchell 2005, unpublished data).
Thescientific interest for peatlands dwellingtestate
amoebae is increasing because they are good indi-
cators of changing environmental conditions. They
are thus used in both ecological (Charman & Warner
1992, Booth 2001) as well as palaeoecological studies
(Schoning et al. 2005, Charman et al. 2006). A more
recent application is their use in applied ecological
researchonhuman impacted peatlands and the history
of land-use changes (Sjögren & Lamentowicz 2006)
or the monitoring of peatland restoration (Davis &
Wilkinson2004).Statisticalmodels commonlyusedin
palaeolimnological research (transfer functions such
as weighted averaging) (Birks 1998) are applied on
the basis of testate amoebae data (Payne et al. 2006).
Testate amoebae are reliable indicators of palaeo-
hydrological conditions in mires. However, can also
indicate pH fluctuations (Patterson et al. 2002).
In this paper we present brief overviews of: (a) the
ecological study of peatland testate amoebae in north-
ern Poland,(b) applications of testateamoeba analysis
in palaeoecological reconstructions.
2 BIOLOGY OF TESTACEA
Testate amoebaeareprotists withacomplex taxonomy.
Theyare regarded as at least biphyleticandaredivided
into Cercozoa (e.g. Euglypha – filose taxa) and Ame-
bozoa (e.g. Difflugia – lobose taxa) (Cavalier-Smith
1997, Foissner 1999,Adl et al. 2005, Nikolaeva et al.
85
2005). Their shared characteristic is the presence of
an external shell, called test from which pseudopods
emerge. There are about 1000 to 2000 described taxa.
The morphology of the test is very important
for identification. Some species produce autogenous
shells, which can be made of pseudo-chitin e.g.:
Archerella (syn. Amphitrema) flavum or Arcella vul-
garis or siliceous plates (called idiosomes) (e.g. Eug-
lypha,Assulina, etc.). Other taxa use foreign material
(referred to as xenosomes) either mineral grains,
diatoms, or other organic material such as fungal
hyphae to build in the test. Some species also form
characteristichornsorotherappendicesthatare useful
for identification e.g. Difflugia leidyi. These different
typesofshells maydifferalso in terms of preservation,
a fact that has implications for palaeoecologists. Thus
tests made of idiosomes are less durable than those
madeof xenosomes,whichinturnareless durablethan
the pseudo-chitin type. But there are some exceptions
e.g. Assulina muscorum, a species that builds shells
from idiosomes, but is extremely resistant and even
observed in pollen preparations (Charman 2001).
Testate amoebae form cysts tosurviveunfavourable
conditions. This ability, in addition to the presence
of a shell allows them to colonize even relatively dry
environments. Short generation period allows them to
respond fast to environmental changes. For example
Centropyxis aculeata was observed to have over 60
generations during the year (Schönborn 1981).
Testate amoebae occur in a broad range of habitats:
marine sand, soil, sea, the bark of trees, and fresh-
water environments. They are usually considered to
be mostly cosmopolitan organisms because they may
disperse easily while encysted in atmospheric dust or
carried passively by migratory birds (Charman et al.
2000). Thus the difference in species composition is
not large between distant regions e.g. in New Zealand
and Great Britain. This situation allows direct com-
parison of species assemblages and interpretation of
differencesintermsofecology. However,somespecies
have restricted Laurasian or Gondwanan distribution,
e.g., Nebela (Apodera) vas (Foissner 1987).
3 STUDY AREA
In2002–2004wecarried out a study on the ecology of
peatland testate amoebae. Our study was restricted to
Sphagnum peatlands, mainly three kettle-hole mires
(Jelenia Wyspa, Jeziorka Kozie and Okoniny) situ-
ated in northern Poland in Tuchola Pinewoods area.
We then used this data in two palaeoecological studies
of Jelenia Wyspa and Tuchola mire.The study area is
presented in Figure 1.
Small lake basins (or mires in this case) are able
to record climatic signals (Moore 2002). We therefore
hypothesized that Tuchola kettle-hole bog also might
have been sensitive to climatic conditions.
Figure 1. Location of the study sites. Description of the
map,surfacesampling mires: 1–Okoniny (OK),2 – Jeziorka
Kozie (KOZ), 3 – Jelenia Wyspa (JEL); coring sites: I –
Tuchola mire, II – Jelenia Wyspa mire; A – town, B – vil-
lage, C – surface waters, D – rail, E – forestry managing
head office.
Polish data are exceptionally important because
Poland is under several contrasted climatic influ-
ences – from oceanic to continental.
Currently we are expanding our focus on Baltic
raised bogs (Kartuzy Lakeland) as well as central
Poland (Łódz region) to address the possible biogeo-
graphic variabilityoftestate amoebae populations and
possible response to climatic gradients.
4 METHODS
Surface sampling was conducted on Sphagnum mires
with the aim describing the ecology of testate amoe-
bae.Foreachsamplingsitemeasurements ofpH,depth
to water table and conductivity were taken. These
parameterswereusedfor descriptionoftestateamoeba
community structure in relation to the environment
Lamentowicz & Mitchell (2005).
Peat cores and monoliths were taken from the
two peatlands for palaeoecological analyses with a
Russian sampler and subsampled in the laboratory.
Pollen, spores and plant macrofossils were also anal-
ysedalongwithtestate amoebae.Thecoresweredated
with 14C method and were fit into the calendar time
scale.
Species-environment correlations were quantified
bymeans ofmultivariatestatistics.A redundancy anal-
ysis (RDA) was used for explaining species relations
86
to environmental parameters (DWT, pH and conduc-
tivity) (Lamentowicz & Mitchell 2005).
Transfer functions to be used in palaecology for
quantitativereconstructionofpastenvironmentalcon-
ditionswereproducedusingeitherweighted averaging
(WA) or weighted averaging partial least squares
(WA-PLS) methods (Birks 1995, Birks 1998). The
optimisation of the transfer function in an ongoing
effort,with newsurface samplesbeingadded andother
improvements to the models (Lamentowicz et al. in
prep).
The redundancy analysis was performed using the
CANOCO software (ter Braak & Šmilauer 1998)
and transfer function inference models were produced
using the software C2 (Juggins 2003). The applica-
tion of these numerical tools allows the integration of
ecological and palaeoecological data sets.
5 RESULTS
5.1 Ecological studies: Species – environment
relations and transfer function models
Atotalof52testateamoebaetaxawererecordedduring
this study. In the redundancy analysis, DWT and pH
explained 20.1% of the variation in the species data.
This analysis allowed us to identify three groups of
taxa: species those are associated with: (1) high DWT
andlowpH, (2)lowDWTand low pH and (3) high pH
and mid-range DWT (Fig. 2A & B).
Thenext stepwastobuildtransferfunctions forspe-
cific environmental variables. Preliminary analyses
revealedthat the simplest model –WA (weightedaver-
aging) performed best for DWT parameter, whereas
WA-PLS model performed best for pH. The perfor-
mance of these models was tested using the surface
samples. The results are in agreement with data from
the other regions of the world (Charman & Warner
1997; Mitchell et al. 1999; Booth 2001).
Despite having gained information on the ecolog-
ical preferences of most taxa, some species, such as
Difflugia urceolata and D. pulex, are still missing
from our training set. This problem exists also in
western European data sets. One possible solution to
this problem is to sample a broader range of ecologi-
cal situations in peatland habitats, e.g. rich fens. The
effort is now concentrated on improving the transfer
functions for palaeoecological reconstructions. Hav-
inggood data abouttheecologyoftestateamoebae,we
can now use them in palaeoecological reconstructions
and monitoring practice.
Until now (December 2005), 76 peatland taxa
were described for Poland, whereas 68 species were
recordedin the modern surfacesamplesand64 species
from the peat material (Lamentowicz, unpubl.).
A selection of indicator testate amoebae taxa are
presented in Figure 3.
Figure 2. Redundancy analysis biplot for species (A) and
samples (B) data. The symbols following the species names
indicatethepercentageofvarianceof each species explained
bythe model(noindication: <15%, *:15–25%, **: 25–45%,
***: 45–66%). Redrawn from Lamentowicz & Mitchell
(2005) with kind permission of Springer Science and Busi-
ness Media.
5.2 Palaeoecological studies: Proxies,
reconstruction, climate, human impact
The core from Tuchola mire yielded a 9000 years
record of the developmental history. Testate amoebae
remains were rare in the early, lake and fen phases of
the development.
87
Figure 3. Pictures of testate amoebae species: high water
levelindicators (A–D), and lowwater levelindicators(E–H);
A–Amphitrema flavum Carter, 1864; B – Arcella dis-
coides Ehrenberg, 1872; C – Arcella gibbosa Penard, 1902;
D–Amphitrema wrightianum Archer, 1869; E – Assulina
muscorum Greeff, 1888; F – Arcella catinus Penard 1890;
G–Difflugia pulex type Penard; H – Euglypha strigosa
Ehrenberg 1872.
Palaeoecologicalreconstruction showedthathydro-
logical conditions were unstable throughout the mire
history. Despite local catchment influences this small
peatland appears to have responded to regional cli-
matic changes. Palynological studies in this area
showed that intensive human impact began in Tuchola
Pinewoods in historical time (Miotk-Szpiganowicz
1992, Obremska & Lamentowicz 2002). This mire
thus contained a mixture of allogenic (climatic) and
anthropogenic (e.g. forest clearance) signals, and the
multiproxy approach proved useful to separate these
two influences.
One species Difflugia urceolata occurred infre-
quently in the development history of the mire. It was
interpreted as an indicator offlooding and eutrophica-
tion of mire (Laminger et al. 1979, Schönborn 1981).
Today, Tuchola mire is flooded every year mainly
during the spring and this situation can be analo-
gousto past conditions where, during moister periods,
flooding was more frequent.
Using the water table depth curve wet shifts in the
history of the mire were recognized and compared
withhydrologicaleventsin Poland and across Europe.
The results of this study will be presented separately
(Lamentowicz et al. in prep.).
Jelenia Wyspa peat archive recorded ca. 2000 years
of local environmental changes. The data shows an
increasing human impact on the mire. It is very dif-
ficult to separate the anthropogenic influences from
climatic ones in this mire. Correlation of results from
three analyses: testate amoebae, palynological and
plant macrofossils show how important the impact
of land-use changes was on the mire. The develop-
ment of Sphagnum mire was most likely induced by
forest clearance approx. 200 years ago. Implementa-
tion of pine monocultures changed considerably the
water chemistry of the mire and caused significant
water table fluctuations. Jelenia Wyspa is a very good
example of a process previously observed also in
other parts of Poland (e.g. Wielkopolska region). The
results of this study will also be presented separately
(Lamentowicz et al. in prep.).
In both of these studies, testate amoebae allowed
quantitative reconstruction of hydrological changes
that would not have been possible with other
palaeoecological methods. The integration of sev-
eral approaches allowed to reaching an integrated
understanding of history of these two sites.
6 CONCLUSIONS
We provided new data on the ecology of testate amoe-
bae in Poland. The data was used to produce a local
transfer function (DWT & pH), filling an important
geographical gap in central Europe. Testate amoebae
were used, for quantitative reconstruction of water
table and pH changes in Polish kettle-hole mires.Tes-
tate amoeba analyses have been used together with
other proxies (pollen and macrofossils). Such multi-
proxy approach is crucial for reconstructing changes
in ecosystems more precisely. Fast-reacting proxies,
e.g. stable isotopes, insects, or aquatic organisms can
extract climatic signal more directly than long-lived
proxiessuchasvegetation(Lotter2003).Testateamoe-
baecan be considered as fast-reactingproxiesthat can
trackabrupt environmentalchangescausedbyclimate
or human.
The results show that palaeomoisture data inferred
fromtestateamoebae can be correlated withother sig-
nals. In Jelenia Wyspa mire, deforestation around the
mire caused a rise and fluctuations of the water table
aswellasanacidification ofthe surface. Tucholamire
peat archive covers a longer time period and is most
likelyinfluenced primarily by climatic changes during
88
the second half of the Holocene. Currently, further
work is carried out on kettle-hole mires and raised
bogs of Pomerania and Central Poland.
Inview oftheongoingand future changeinclimate,
we need further high-resolution palaeoclimatological
studies in order to gain a better understanding of
the response of natural ecosystems to past climatic
changes. Testate amoebae are very valuable indica-
tors for studying the relationships between peatlands,
climate, and human activities (Charman 2001).
In addition to classical palaeoecological studies,
testate amoebae can be employed to monitoring mire
restoration(Davis&Wilkinson 2004), or spontaneous
regeneration (Chapman et al. 2003).
By combining palaeoecological and ecological
information we can answer the question – to which
conditions we want to restore a site? Knowledge
about the past of peatlands provides vital informa-
tion for managers. We therefore call for including
paleoecologyin the classical toolbox of peatland man-
agement. Having sound knowledge on the develop-
mental history of the peatlands that we restore is not a
luxury!
ACKNOWLEDGEMENTS
This paper was completed as the result of research
projects funded by the Polish Ministry of Scientific
Research and Information Technology: “Development
andoriginof miressituatedinSta˛˙zkaValleyinTuchola
Landscape Park” No. 3P04G04323 (Principal Inves-
tigator: Mariusz Lamentowicz). The training set has
been developing during the realization of the sci-
entific grant No. 2P04G03228 Climatic changes in
Pomerania (N Poland) in the last millennium based
on the multiproxy high-resolution studies funded by
the Polish Ministry of Scientific Research and Infor-
mation Technology (Principal Investigator – Mariusz
Lamentowicz). Edward Mitchell is supported by EU
project RECIPE (Reconciling Commercial Exploita-
tion of Peat with Biodiversity in Peatlands Ecosys-
tems). RECIPE is partly supported by the European
Commission (n◦. EVK2-2002-00269) and partly, for
the Swiss partners EPFL and WSL-AR, by the OFES
(Swiss Federal Office for Education and Science),
Switzerland.
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