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Phytolith assemblages as indicators ofcoastal environmental changes andhurricane overwash depositionHou-Yuan Lu1,2 and Kam-biu Liu2*
(1Institute of Geology and Geophysics, Chinese Academy of Sciences, P.O. Box
9825, Beijing 100029, China; 2Department of Geography and Anthropology,Louisiana State University, 227 Howe-Russell Geoscience Complex, Baton RougeLA 70803, USA)
Received 1 August 2004; revised manuscript accepted 12 January 2005
Abstract: We demonstrate that phytolith assemblages are a useful proxy for reconstructing coastal
environmental changes and for validating the overwash origin of sand layers in palaeotempestology
studies. Phytolith analysis was conducted on 50 topsoil or surface sediment samples collected from a
variety of coastal plant communities or depositional environments in the southeastern USA. The data
suggest that different coastal subenvironments can be distinguished by their modern phytolith assemblages.
For example, coastal marsh samples contain a diverse phytolith assemblage dominated by smooth-elongate
and square morpho-types and maritime forest samples are dominated by phytoliths from palms and
broadleaf dicotyledonous plants. Remarkably, the phytolith assemblages from sand dunes are
characterized by high percentages of two-horned-tower, flat-tower, spool/horned-tower and short-saddle
types. Phytolith analysis of three prehistoric sand layers in a sediment core from Western Lake,
northwestern Florida, shows that they contain a phytolith assemblage similar to those characteristic of
sand dunes and interdune meadows. These observations are confirmed by the results of principal
components analysis and discriminant analysis on the modern and fossil phytolith data sets. Our study
results support the interpretation that the sand layers in Western Lake were indeed formed by the erosion
of sand dunes during overwash processes caused by landfalling catastrophic hurricanes.
Key words: Phytoliths, hurricanes, palaeotempestology, coastal environments, sand dunes, overwash
deposition, Atlantic Coast, Gulf of Mexico, late Holocene.
Introduction
In the emerging field of palaeotempestology (Liu and Fearn,
2000a; Liu, 2004), the proxy proven to be most useful in the
reconstruction of past hurricane strikes comes from overwash
sand layers preserved in sediment cores retrieved from coastal
lakes and marshes (Liu and Fearn, 1993, 2000a,b; Zhou, 1998;
Donnelly et al., 2001a,b; Scott et al., 2003; Liu, 2004). The
principle underlying this research methodology is that when an
intense hurricane makes landfall and causes a storm surge that
exceeds the height of the sand barrier, sand will be washed over
from the beach and dunes into the backbarrier lakes and
marshes, forming an overwash fan (Liu, 2004). Although sand
layers of recent age (e.g., twentieth century) found in some
coastal sediment cores can be reliably attributed to historic
storm events and their associated overwash processes by
examining historic aerial photos of coastal sand transport
and by means of radiometric (137Cs and 210Pb) dating of these
cores (Donnelly et al., 2001a,b), independent evidence is
needed to verify the overwash origin of the older, especially
prehistoric, sand layers because the sand could have come from
other coastal or even fluvial environments rather than the sand
dunes or beach ridges.
Microfossils such as dinoflagellates, foraminifera, diatoms
and pollen have been used as indicators of saltwater intrusion
into coastal lakes and marshes resulting from past storm surges
and tsunamis (Hemphill-Haley, 1996; Zhou, 1998; Collins
et al., 1999; Hippensteel and Martin, 1999; Liu and Fearn,
2000b; Hughes et al., 2002; Liu et al., 2003; Scott et al., 2003).
However, some of these conventional proxies for environmen-
tal reconstruction, such as pollen, are not very useful for
deciphering the provenance of the sand in the sand layers and
for identifying the coastal sandy subenvironments from which
the sand is derived. In this paper we demonstrate that modern
phytolith assemblages in soil or sediment samples can be used
to differentiate different coastal subenvironments such as sand*Author for correspondence (e-mail: kliu1@lsu.edu)
The Holocene 15,7 (2005) pp. 965�/972
# 2005 Edward Arnold (Publishers) Ltd 10.1191/0959683605hl870ra
dunes, marshes, freshwater lakes, estuarine and bay bottoms
and maritime forests. In particular, phytolith analysis can be a
useful tool to test the hypothesis that the sand in the sand
layers used in palaeotempestology studies does indeed come
from the coastal sand dunes resulting from overwash processes.
Material and methods
Phytolith surface samplesFifty surface samples were collected for phytolith analysis from
three coastal locations in the southeastern USA (Figure 1A).
Cumberland Island (Figure 1B), a barrier island in southern
Georgia, is fringed by a well-developed continuous line of
beach and high dunes along the Atlantic coast. Interspersed
between and behind the beach and sand dunes is a complex
array of freshwater lakes, small Juncus/sedge fens, interdune
meadows and upland maritime forests dominated by various
species of oaks, pines, palms and other hardwoods.
Atchafalaya Marsh (Figure 1C), southern Louisiana, is a
large coastal/estuarine marsh that extends along the mouth of
the Atchafalaya River and the west shore of Atchafalaya Bay
to the Gulf of Mexico. This large coastal marsh exhibits a
salinity gradient from cypress swamp, fresh marsh, brackish
marsh, to salt marsh, whereas the Atchafalaya Bay itself offers
a distinct set of open-water, estuarine bay depositional
environments.
Western Lake (Figure 1D), a backbarrier oligohaline lake
within Grayton Beach State Park, northwestern Florida, is
situated behind a barrier beach about 150�/200 m wide and a
continuous line of sand dunes rising to 6.2�/9.3 m above sea
level (Liu and Fearn, 2000a). Vegetation around Western Lake
is a typical Gulf coastal community (Moreno-Casasola, 1988)
consisting of sea oats (Uniola paniculata) growing on the sand
dunes, cordgrass (Spartina alterniflora) growing on the seaward
edge of the dunes and marshes around the lake and pine
flatwoods, scrub oak thickets and mixed hardwood-palmetto
woodlands occurring on the uplands and hammocks.
The 50 surface samples (Figure 1) were collected from nine
different coastal vegetation communities or depositional en-
vironments representing different ecological conditions: fresh
marsh, brackish marsh, salt marsh, maritime forest, freshwater
lake, Juncus/sedge fen, interdune meadow, primary dune and
estuarine bay. Terrestrial samples were collected from the
uppermost 2�/3 cm of the topsoil (forest, marsh and dune)
by means of a small spade. Those derived from the surface mud
(lakes and estuarine bay) were collected by means of an
Ekman’s dredge.
The samples were prepared by a procedure slightly modified
from Piperno (1988) and Runge (1999). It consists of sodium
pyrophosphate (Na4P2O7) deflocculation, treatment with 30%
hydrogen peroxide (H2O2) and cold 15% hydrochloric acid
(HCl), zinc bromide (ZnBr2, density 2.35 g/cm3) heavy liquid
separation and mounting on a slide with Canada Balsam.
Phytolith counting and identification was performed using a
Nikon Optiphot microscope at 400�/ magnification. More
than 350 phytolith grains were counted in each sample.
Identification was aided by the use of reference materials
(Lu and Liu, 2003a,b) and published keys (Piperno, 1988;
Mulholland and Rapp, 1992, Kondo et al., 1994; Piperno and
Pearsall, 1998; Runge, 1999, Pearsall, 2000).
Sediment cores from Western LakeTo test if the modern phytolith data can be used to aid the
reconstruction of coastal palaeoenvironmental changes and the
identification of the provenance of lake sediments, we analysed
the phytolith contents of a short core (core 9) from Western
Figure 1 A. Location of Cumberland Island (Georgia), Atchafalaya Marsh (Louisiana) and Western Lake (Florida) on the Atlantic coastand Gulf coast of southeastern USA. B. Major habitat types and location of surface samples (dots) (C27-47) on Cumberland Island. C.Marsh salinity zones and location of surface samples (A1�/26) in Atchafalaya Marsh. D. Geomorphic setting of Western Lake and locationof surface samples (W48-50) and core 1 and core 9 (triangles). Contours are in metres
966 The Holocene 15 (2005)
Lake, northwestern Florida (Figure 1). A 6.5-m core (core 1)
from Western Lake has yielded a 5000-yr proxy record of
hurricane activity that includes 12 direct strikes by catastrophic
hurricanes over the past 3400 years (Liu and Fearn, 2000a).
The same processing method was used on the core samples as
on the surface samples for the extraction of phytoliths.
Statistical analysisBy using the statistical program CANOCO 4.0 (ter Braak,
1996), principal components analysis (PCA) was performed on
the 50 surface samples to reveal the internal structure of the
modern phytolith data set. The PCA results provided a basis to
classify the surface samples into statistically meaningful
groups, which could be interpreted in terms of the coastal
vegetation communities or depositional environments from
which the surface samples were collected. In addition, dis-
criminant analysis (sensu Liu and Lam, 1985), an inferential
statistical technique, was performed using SPSS version 10.0 to
classify the 50 phytolith samples with reference to the a
priori groups suggested by the PCA results. The discriminant
functions derived from these surface samples were then
applied to the fossil phytolith data obtained from the Western
Lake core to predict the group membership of each fossil
phytolith sample. These predicted group memberships repre-
sent the palaeovegetation communities from which the fossil
phytolith assemblages are derived and therefore shed light
on the provenance of the sediment that contain them
(Liu and Lam, 1985).
Results
Phytolith assemblages in modern coastalsubenvironmentsThirty phytolith morpho-types (including two ‘unknown’
morpho-types) were recognized in the 50 surface samples
(Figure 2). Figure 3 shows some of the more common or
characteristic types. All surface samples from coastal marshes
are characterized by high percentages of smooth-elongate
(169/6%) (Figure 3j) and square (119/6%) (Figure 3n)
phytoliths. Remarkably, marshes of different salinity environ-
ments seem to be distinguishable by the occurrence of certain
associated phytolith types. Fresh marsh samples tend to have
higher percentages of dumbbell (Figure 3a) and sinuate-
elongate (Figure 3i) phytolith types than the brackish and
salt marshes and the presence of Pteridophyte phytoliths
reflects the presence of ferns in fresh marshes. Between the
brackish marsh and salt marsh samples, brackish marsh
samples seem to have higher percentages of spool/horned
towers (Figure 3d), whereas salt marsh samples generally
have higher percentages of fan-shaped (79/5%) and Spartina-
type (69/2%) phytoliths. The Spartina-type (Figure 3f) is
a rondel/saddle ellipsoid morpho-type typically produced
by Spartina alterniflora, the dominant grass of salt marshes
(Lu and Liu, 2003b).
Phytolith assemblages from the topsoil samples of maritime
forests on Cumberland Island are characterized by very high
percentages of broadleaf-types (309/23%) (Figure 3l, m) and
Palmae-type 1 (439/28%) (Figure 3g). The broadleaf-types
include many phytolith morpho-types (Piperno, 1988; Kondo
et al., 1994; Runge, 1999) such as the elongate phytolith type
A4 (equivalent to broadleaf-type 1 in this study, Figure 3l) and
type A5 (equivalent to broadleaf-type 2 in this study, Figure
3m) of Runge (1999). They are characterized by the presence of
facets and are mainly produced by broadleaf trees or shrubs.
Palmae-type 1 (Figure 3g) is a spherical phytolith with a
spinulose surface, typically produced by palms (e.g., Piperno,
1988; Runge, 1999).
Phytolith assemblages from freshwater coastal lakes on
Cumberland Island are different from those of the adjacent
maritime forests in having very high percentages of both
Palmae-type 1 (749/6%) and Palmae-type 2 (99/6%), whereas
the broadleaf types are absent. Palmae-type 2 is a spherical
Figure 2 Diagram of phytolith percentages from 50 surface samples from Cumberland (C), Atchafalaya Marsh (A) and Western lake (W)arranged by coastal subenvironments from which they were collected
Hou-Yuan Lu and Kam-biu Liu: Phytolith assemblages in coastal palaeotempestology 967
phytolith with a regularly spinulose surface like Palmae-type 1,
but the spinule apices are round instead of sharp; they are
probably derived from a specific member of the Palmae family
or from another monocotyledonous plant (Runge, 1999).
Three surface samples from Juncus/sedge fens in Cumber-
land Island are characterized by abundant sinuate-elongate,
smooth-elongate and Cyperaceae type phytoliths (8.0�/12.1%).
The abundance of the Cyperaceae type is attributed to the
predominance of sedges (especially Carex spp.) in these small
wetlands, some of which are probably developed on dried or
ephemeral lake beds. The spikes of broadleaf-type 2 and
Palmae-type 1 in two samples collected near the edges of these
small fens are probably due to local influence from the adjacent
forest vegetation.
The phytolith composition of the surface samples from
interdune meadows shows very high frequencies of dumbbell
phytoliths (479/13%). Cross-shaped phytoliths (Figure 3b)
reach values of around 2�/8%. Dumbbell and cross phytoliths
Figure 3 Photomicrographs of the main phytolith morpho-types counted in this study: (a) dumbbell, (b) cross, (c) long-saddle, (d) spool/horned tower, (e) flat tower and two-horned tower, (f) Spartina-type, (g) palmae-type 1, (h) short-saddle, (i) sinuate-elongate, (j) smooth-elongate, (k) long-point, (l) broadleaf-type 1, (m) broadleaf-type 2, (n) square
968 The Holocene 15 (2005)
are produced at high percentages by grasses of the Panicoideae
subfamily, such as Cenchrus incertus and Panicum hemitomon
(Lu and Liu, 2003b).
The modern phytolith composition of the primary dunes in
both Cumberland Island and Western Lake is characterized by
high amounts of two-horned-tower (309/25%) (Figure 3e), flat-
tower (139/7%) (Figure 3e), spool/horned-tower (149/6%) and
short-saddle (99/6%) (Figure 3h) phytoliths. This assemblage is
produced by the dominant grasses of coastal sand dunes, such
as Uniola paniculata and Aristida desmantha (Lu and Liu,
2003b). Although, to some extent, these characteristics are also
found in samples from the coastal marshes, the dune samples
can be distinguished from the marsh samples by having very
low percentages of square, smooth-elongate, sinuate-elongate
and long-point types (Figure 3k).
The phytolith composition in the surface samples of
estuarine bay bottoms is characterized by high percentages of
fan-shaped, square, smooth-elongate and long-point phytoliths
of silt sizes (109/7%, 139/9%, 289/7%, 89/4%, respectively)
that are derived from epidermal long-cells, trichome-cells and
bulliform-cells of grasses. Not surprisingly, these samples from
the Atchafalaya Bay are generally similar to the marsh samples
collected from the adjacent Atchafalaya Marsh in terms of
their predominant morpho-types. However, the estuarine bay
bottom samples can be distinguished from the marsh samples
by having no or few dumbbell, spool/horned-tower and
Spartina-type phytoliths. The lower diversity of phytoliths in
the estuarine and bay bottom samples may be due to
differential dispersal and deposition abilities of different
phytolith morpho-types in a fluvial-estuarine environment.
Numerical analysis of modern phytolith dataPercentage data from 23 phytolith types (see Figure 4a) were
used in the principal components analysis (PCA) and dis-
criminant analysis. The PCA biplots of the 23 phytolith types
and the 50 surface samples are shown in Figure 4. The first two
principal components, axis 1 and axis 2, account for 56.1% and
18.8% of the variance in the phytolith data, respectively. On the
biplot of principal component loadings for the 23 phytolith
types (Figure 4a), Palmae-types and broadleaf-type 1 have the
highest loadings on axis 1, whereas two-horned-tower, flat-
tower and dumbbell�/cross have the highest loadings on axis 2.
On the biplot for principal component scores for the 50 surface
samples (Figure 4b), samples from freshwater lake and
maritime forest sites have the highest positive scores on axis
1, where the Palmae-types and broadleaf-type 1 also have the
highest loadings. Samples from estuarine bay and marsh sites
score most negatively on the far left of axis 1, corresponding to
highly negative loadings for fan-shaped, square, smooth-
elongate and long-point phytolith types. The second principal
component, axis 2, primarily distinguishes the samples from
sand dunes and interdune meadows from the rest of the
samples. Thus, the results of the PCA ordination analysis
suggest that the 50 surface samples can be divided into five
primary groups: (1) estuarine bay; (2) marsh (including fresh
marsh, brackish marsh and salt marsh); (3) Juncus/sedge fen;
(4) sand dune/interdune meadow; and (5) freshwater lake/
maritime forest. With only a few exceptions (e.g., A18, C37,
C46, C47), the phytolith ‘signatures’ of these five groups are
quite well-defined with little overlap among them (Figure 4a,
see also Figure 2).
After these five groups were delineated by PCA, discrimi-
nant analysis was used to validate the classification and to
generate probability estimates for statistical inference (Liu and
Lam, 1985). All (100%) of the samples were correctly classified
into their respective a priori groups (Table 1). Figure 5 shows
the five groups of surface samples plotted against discriminant
functions 1 and 2, which account for 40.1% and 33.5% of the
variance, respectively. Again, the five groups and their cen-
troids are clearly distinct from each other with very little
overlap between groups. The high degree of correct classifica-
tion suggests that the classification of the surface samples into
the five groups is statistically robust.
These surface sample results indicate that the phytoliths
deposited in the different coastal subenvironments in our study
areas are reliable indicators of the major vegetation commu-
nities at or around the site. These results can then be applied to
the fossil phytolith spectra from a sediment core from Western
Lake to reconstruct palaeoenvironmental changes and to infer
the provenance of the sediment.
Figure 4 a. PCA biplot showing coordinates of the 23 phytolithtypes (vectors) plotted along axis 1 and axis 2. Abbreviationfor phytolith types: Broad1, broadleaf-type 1; Broad2, broadleaf-type 2; Cypera, cyperaceae type; Dumb�/Cross, dumbbell andcross; Fan, fan-shaped; Ftower, flat-tower; Gymnos, gymnospermtype; LPoint, long-point; LSaddl, long-saddle; Palma1, palmae-type 1; Palma2, palmae-type 2; Pterid, pteridophyte type; Rectan,rectangle; RFan, fan-reed; Rondel, rondel; Selong, sinuate-elon-gate; SHTower, spool/horned tower; Smlong, smooth-elongate;Sparti, Spartina type; SPoint, short-point; Square, square; SSaddl,short-saddle; THTower, two-horned tower. b. PCA biplot showingcoordinates of the 50 surface samples plotted along axis 1 andaxis 2. Serial numbers of the samples are the same as in Figures 1and 2
Hou-Yuan Lu and Kam-biu Liu: Phytolith assemblages in coastal palaeotempestology 969
Phytolith record from Western LakeCore 9 from Western Lake, a short core taken near core 1,
contains 90 cm of organic lake mud (organic matter content
10�/13% as determined by loss-on-ignition) interrupted by
seven sand layers. These sand layers have been interpreted to be
formed by overwash processes occurring during strikes by
catastrophic hurricanes of category 4 or 5 intensity according
to the Saffir/Simpson scale (Liu and Fearn, 2000a). For the
phytolith analysis we selected eleven samples from the organic
lake mud and three samples from three thick sand layers
(referred to as sand layers 1, 2 and 3, respectively) occurring at
depths of 52.0�/53.0 cm, 58.5�/60.0 cm and 66.0�/68.5 cm as
measured on the core (Figure 6). These three sand layers
were selected because they were sufficiently thick to provide
clean samples of sand uncontaminated by the embedding
organic sediments. These prominent sand layers were radio-
carbon dated to 13209/50, 14109/50 and 18509/50 yr BP
(uncalibrated 14C ages), respectively (Liu and Fearn, 2000a).
Phytolith analysis of the eleven organic lake sediment
samples shows that they are dominated by the Palmae-type 2
(419/9%), Palmae-type 1 (129/3%) and smooth-elongate (9.69/
3.7%) morpho-types. Dumbbell and cross (7.49/3.9%) and
flat-tower and two-horned-tower (5.89/2.5%) morpho-types
are also present consistently in the organic mud at lower
percentages (Figure 6). The presence of broadleaf-type 1 and
broadleaf-type 2 phytoliths is also notable. This apparently
mixed assemblage suggests that the organic sediments in
Western Lake come from multiple sources, but the principal
contributors are maritime forests and freshwater wetlands
around the lake. On the other hand, the sand sampled from
the three thick sand layers contains very high percentages of
flat-tower phytoliths (at 20.1%, 18.6% and 51.1% in sand layers
1, 2 and 3, respectively), along with somewhat elevated
percentages of short-saddle and two-horned-tower phytoliths.
This unusual assemblage, especially the prominent spikes of the
flat-tower type, is similar to the modern phytolith assemblages
characteristic of the sand dunes near Western Lake and in
Cumberland Island.
The contrast in phytolith assemblages between the organic
lake sediment and the sand layers is confirmed by the results
of discriminant analysis. The same discriminant functions
derived from the surface sample data set were used to classify
the 14 fossil phytolith samples. All 11 samples from organic
lake sediment were classified as freshwater lake/maritime
forest (group 4), whereas the three samples from the sand
layers were classified as sand dune/interdune meadow (group 5)
(Table 1; Figure 6). When the fossil samples from Western
Lake are plotted against the same discriminant functions as
the surface samples (Figure 5), the three sand layer samples
are located closer to the centroid of the sand dune/interdune
meadow group than the rest of the core samples. Therefore,
the phytolith evidence from the Western Lake core suggests
that the sand in the three prominant sand layers was derived
from the sand dunes separating the lake and the Gulf. Grasses
growing on the dune ridges, such as Uniola paniculata,
Aristida desmantha and Panicum amarum, are the principal
producers of flat-tower and two-horned-tower phytoliths
Table 1 Classification results of the discriminant analysis performed on the 50 surface samples from the five vegetation groups and on 14fossil samples (bold) from core 9 of Western lake
Group no. Actual group Predicted group membership Total cases
Estuarine bay Marsh Juncus/
sedge fen
Sand dune/
interdune meadow
Freshwater lake/
maritime forest
Group 1 Group 2 Group 3 Group 4 Group 5
1 Estuarine Bay 14a (100%)b 0 0 0 0 14
2 Marsh 0 14 (100%) 0 0 0 14
3 Juncus/sedge fen 0 0 3 (100%) 0 0 03
4 Sand dune/interdune meadow 0 0 0 8 (100%) 0 8
5 Freshwater lake/ Maritime forest 0 0 11 (100%) 11
Ungrouped (Core 9 samples) 0 0 0 3 (21.4%) 11 (78.6%) 14
aNumber of samples classified as that group.bPercentage of samples classified as that group.
Figure 5 The 50 surface samples plotted against canonicaldiscriminant functions 1 and 2 and their classification into fivegroups corresponding to five vegetation communities. Also plottedare the 14 fossil samples from Western Lake core 9. Samples fromthe three prominent sand layers interpreted to be of overwashorigin are labelled as sand 1, 2 and 3
970 The Holocene 15 (2005)
(Lu and Liu, 2003b). Overwash processes associated with
catastrophic hurricane landfalls are the most likely mechanism
that washed sand from the sand dunes to the bottom of
Western Lake.
Discussion and conclusions
Previous palaeoecological applications of phytolith studies
have focused mainly on the reconstructions of broad vegetation
types at regional or continental scales, such as temperate or
tropical grasslands (e.g., Fredlund and Tieszen, 1994; Barboni
et al., 1999; Blinnikov et al., 2002) as well as Mediterranean
woodlands and tropical forests (Piperno and Becker, 1996;
Delhon et al., 2003; Piperno and Jones, 2003). Except in a few
cases (Fearn, 1998; Horrocks et al., 2000), phytolith analysis
has not been applied to the study of coastal environmental
changes, especially at the finer spatial scales that identify
coastal subenvironments such as salt marshes, maritime forests
and sand dunes.
In this paper, we demonstrate that different coastal sub-
environments can be distinguished by their modern phytolith
assemblages. Notably, surface samples from coastal marshes
contain a diverse phytolith assemblage dominated by the
smooth-elongate and square types, whereas soil samples from
maritime forests are dominated by phytoliths from palms and
broadleaf dicotyledonous plants. Moreover, surface samples
from coastal sand dunes are characterized by high percentages
of two-horned-tower, flat-tower, spool/horned-tower and
short-saddle phytoliths. These results confirm and reinforce
our previous conclusion that grasses typically growing on sand
dunes (e.g., Uniola paniculata, Aristida desmantha) produce
distinctive phytolith assemblages (Lu and Liu, 2003b). Thus,
phytoliths found in the sand layers of sediment cores taken
from Western Lake could be used to reveal the provenance of
the sand that embeds them. Data from core 9 show that the
three most prominent sand layers contain a phytolith assem-
blage similar to those derived from the grass plants and
sediment samples from the primary dunes. This finding
supports our previous interpretation that the sand layers
were formed by the erosion of sand dunes during overwash
processes caused by catastrophic hurricane landfalls (Liu and
Fearn, 2000a). This study demonstrates that, in association
with overwash sand layers, phytoliths can be used as a useful
proxy for the reconstruction of past hurricane activity from
coastal lake sediments.
Acknowledgements
We thank J. Blackburn, D. Cahoon, M. Fearn, S. Fournet,
C.A. Reese, C.M. Shen, Z. Yao and X.Y. Zhou for assistance
with fieldwork and M. Eggart for cartographic assistance. This
work was supported by NSFC (40024202, 40325002), the Risk
Prediction Initiative (RPI) of the Bermuda Biological Station
for Research (RPI-00-1-002) and the US National Science
Foundation (SES-9122058; BCS-0213884).
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Figure 6 Percentage diagram of major phytolith types in core 9 of from Western Lake. Phytolith spectra from the three prominent sandlayers are delimited by dotted lines. Radiocarbon dates are shown on the right
Hou-Yuan Lu and Kam-biu Liu: Phytolith assemblages in coastal palaeotempestology 971
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