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Institute of Geology and Geophysics, Chinese Academy of Sciences, P.O. Box 9825,
Beijing 100029, China
b
Department of Geography and Anthropology, Louisiana State University, Baton Rouge, LA 70803, USA
Received 28 October 2002; received in revised form 8 May 2003; accepted 8 May 2003
Abstract
Thirty-four grass species were collected for phytolith analysis from a variety of coastal environments in the southeastern USA
(Georgia, Florida, and Louisiana), including salt marshes, freshwater/brackish marshes, pine/oak forests, maritime hardwood
forests, and sand dunes. Phytoliths produced by these modern grasses include a large diversity of shapes and types. We propose
a preliminary relationship between modern coastal plant communities and their predominant phytolith contents. The dominant
grasses of coastal sand dunes, such as Uniola paniculata, produce primarily at tower and two-horned tower phytoliths. Rondel/
saddle ellipsoid phytoliths are mainly produced by Spartina alterniora, the most common plant in coastal salt marshes. Rondel and
spool/horned tower phytoliths are common in brackish marsh grasses. Plants from interdune meadow produce primarily dumbbell
phytoliths, as well as small cross and Cyperaceae-type phytoliths. These results provide a basis for the interpretation of fossil
phytolith assemblages and the reconstruction of coastal environmental changes.
2003 Elsevier Ltd. All rights reserved.
Keywords: phytoliths; silica bodies; grasses; microfossils; coastal environments; Quaternary; southeastern USA
1. Introduction
Phytoliths are microscopic silica bodies that precipitate in or between cells of living plant tissues. They
occur in many plant families (Pearsall, 2000; Piperno,
1988, 2001), but are especially abundant, diverse, and
distinctive in the grass family (Gramineae) (Blackman,
1971; Brown, 1984; Cliord & Watson, 1977; Grob,
1896; Piperno & Pearsall, 1998; Prat, 1936; Twiss, 2001).
Many taxa in Gramineae are characterized by phytoliths
with specic morphological characteristics, hence their
taxonomic signicance. Phytoliths are released from
plant tissues when they are decayed, burned, or digested.
Released phytoliths thus become microfossils of the
plants that produce them.
* Corresponding author.
E-mail address: kliu1@lsu.edu (Kam-biu Liu).
0272-7714/03/$ - see front matter 2003 Elsevier Ltd. All rights reserved.
doi:10.1016/S0272-7714(03)00137-9
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2. Phytolith classication
At present, the classication of phytoliths is principally based on the study of some modern Gramineae,
Xylophyta, and a few other plants. Dierent researchers
have suggested dierent terms and classication schemes
because of the dierences in materials, classication
criteria, and study areas. So far, no uniform and
convenient classication scheme has been widely adopted for various conditions. Nevertheless, many important investigations have made a great contribution to
phytolith classication. Taking Gramineae phytoliths as
an example, Prat (1936) divided the Panicoideae subfamily into two groups by studying the shapes of short
cells in the leaf veins of some genera of Gramineae.
Twiss, Suess, and Smith (1969) classied Gramineae
phytoliths into four groups after they summarized the
achievements of Prat (1936) and other researchers.
Brown (1984) classied Gramineae phytoliths into eight
categories including more than 130 types after studying
the phytoliths from dierent parts of 112 taxa of
Gramineae plants. Piperno (1988) presented an index
table of two dierent kinds of phytoliths. Mulholland
and Rapp (1992) proposed one morphological classication of grass silica bodies after summarizing dierent
researchers classications for the Gramineae family.
Piperno and Pearsall (1998) summarized the distribution
of short-cell phytoliths from the grasses, which were
described as circular to oval-(rondels), saddle-, bilobate-,
or cross-shaped, and well-established diagnostic features
of the leaf epidermis.
The above classication of phytoliths was mainly
proposed by European and American botanists. Japanese researchers such as Sase and Kondo (1974)
developed a morphological classication, which added
the following three classes: fan-shape, point-shape, and
Table 1
Comparison of dierent grass phytolith classication systems
Lu et al., 1996 and
Wang and Lu, 1993
Piperno and
Pearsall, 1998
(short cell)
Dumbbell
Brown, 1984
Bilobate
Panicoid class
(dumbbell)
VI Bilobates
(Complex dumbbell)
VII Polylobates?
Cross
(Cross)
VIII Crosses
Chioridoid class
(thin Chloridoid)
IV Saddles
Sase and
Kondo,
1974
Kondo et al.,
1994
Mulholland and
Rapp, 1992
(short cell)
Panicoid shapes
(dumbbell, angular,
nodular, cross,
crenate,
half dumbbell)
Panicoid
Panicoid class
Dumbbell
(Complex regular
bilobate?)
Cross
Chloridoid shapes
(saddles)
Chloridoid
Bambusid class
Collapsed saddle
Saddle: one
concave edge
Saddle: no
concave edges
Narrow-elliptate
Short saddle
Multiple tooth
Weakly tooth
Hat
Fan type with
raised ridge
Fan type without
raised ridge
Square
Rectangle
Plate-like bar
Point (joint) bar
Smooth-bar
Long-point
Short-point
Short saddle
Plateau saddle
Conical?
Bilobate/saddle conical
irregular short cell
Wavy trapezoid
Saddle: two
concave edges
(Chloridoid)
Festucoid class
(circular rectangular
elliptical oblong)
Rondel
(circular to oval)
Elongate class
Chloridoid class
V Trapezoids
Horned towers,
at towers, regular spools
Half rotated, angular,
irregular spool
Festucoid shapes
(square/rectangular)
Pooid
(Festuoid)
Chionochloid class
Triangle?
Phytolith from
other short cell
Festucoid class
(boat-shaped)
Pentagon?
(Round/oblong)
(Hat/cone-shaped)
I Plates (IB)
Bulliform cells
Fan-shaped
Fan-shaped class
I Plates (IA)
Long cells
Elongate
Elongate class
II Trichomes
Trichomes
Point-shaped
Point-shaped class
Sinuate:
polyloate
bilobate?
Rectangle
Rondel (entire,
at, indented)
H. Lu, K.B. Liu / Estuarine, Coastal and Shelf Science 58 (2003) 587600
Long saddle
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Fig. 1. Hand drawings of principal morpho-types of short-cell phytoliths found in the 34 species of coastal grasses from the southeastern USA.
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Table 2
List of 34 modern grasses from the US Atlantic and Gulf coasts used for the phytolith analysisa
No.
Name
Subfamily (Gould
& Shaw, 1983)
1
2
3
4
5
6
7
8
9
10
11
12
13
Panicoideae
Panicoideae
Panicoideae
Chloridoideae
Bambusoideae
Bambusoideae
Pooideae
Panicoideae
Arundinoideae
Arundinoideae
Chloridoideae
Chloridoideae
Chloridoideae
14
15
16
17
18
Chloridoideae
Chloridoideae
Panicoideae
Chloridoideae
Oryzoideae
19
20
Panicoideae
Panicoideae
21
Panicoideae
22
Panicoideae
23
Panicum virgatum L.
Panicoideae
24
25
26
27
28
Arundinoideae
Pooideae
Panicoideae
Panicoideae
Panicoideae
29
30
31
32
33
34
Panicoideae
Chloridoideae
Chloridoideae
Chloridoideae
Chloridoideae
Oryzoideae
All plant samples are preserved in the Department of Geography and Anthropology, Louisiana State University, Baton Rouge, LA 70803, USA.
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Panicoideae
Andropogon glomeratus
Andropogon ternaries
Anthaenatia rufa
Cenchrus incertus
Erianthus strictus
Panicum amarum
Panicum dichotomiorum
Panicum hemitomon
Panicum verrucosum
Panicum virgatum
Saccharum ocinarum
Setaria sp.
Sorgastrum nutans
Sorghum halepense
A*
A*
A*
A*
A*
A*
A*
A*
A*
A*
A*
A*
A*
A*
Chloridoideae
Aristida desmantha
Ctenium aromaticum
Dactyloctenium
aegyptium
Distichlis spicata
Eragrostis oxylepis
Eragrostis cilianensis
Eustachys petraea
Spartina alterniora
Spartina patens
Sporobolus virginicus
Uniola paniculate
C
C*
R*
C*
C*
R
R
R
C
R*
A*
R?
A?
A
C
A*
R
A*
A*
A*
R*
R
Bambusoideae
Arundinaria longifolia
Arundinaria gigantean
A*
A*
Oryzoideae
Leersia oryzoides
Zizaniopsis miliacea
R*
C*
A
A
R
R
R
R
A
C
R
C
A
A
C
R
R
A
R
A*
A*
R
R
R
A*
C
R
R
A
A
C
R
R
C
R
R
R
R
R
Pooideae
Avena sativa
Poa annua
Arundinoideae
Chasmanthium laxum
Chasmanthium
ornithorhynchum
Phragmites australis
C
R
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Plate 1. Illustrations of long-cell and short-cell phytolith morpho-types in four common coastal grass species: A, Panicum verrucosum (mostly
dumbbells); B, Leersia oryzoides (mostly dumbbells, some at towers); C, Aristida desmantha (mostly tower-shaped phytoliths, some dumbbells);
D, Panicum hemitomon (mostly dumbbells). One scale 10 lm.
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595
Plate 2. Illustrations and photomicrographs of dierent phytolith morpho-types in three common coastal grass species: A, Spartina alterniora
(mostly rondel/saddle ellipsoid phytoliths); B, Uniola paniculata (mostly at towers, some two-horned towers and spool/horned towers); and C,
Spartina patens (mostly spool/horned towers).
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Plate 3. Photomicrographs of principal phytolith morpho-types in four common coastal grass species: A, Arundinaria longifolia (mostly long
saddles); B, Avena sativa (mostly wavy trapezoid); C, Dactyloctenium aegyptium (mostly short saddles); and D, Phragmites australis (mostly plateau
saddles) (bar unit in this plate is lm).
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Fig. 2. An idealized transect of coastal environments in the southeastern USA showing major vegetation zones and their representative phytolith
assemblages. The width of a black bar indicates relative abundance of each phytolith type.
5. Conclusions
Phytolith analysis of modern grasses from various
coastal environments including salt marshes, brackish
marshes, freshwater marshes, pine/oak forests, maritime
hardwood forests, and sand dunes in the southeastern
USA has revealed a large diversity of phytolith shapes.
The dominant grasses from coastal sand dunes, such as
Uniola paniculata, and Eragrostis spp., produce primarily at towers and two-horned towers, as well as small
short saddle and dumbbell-shaped phytoliths. The
grasses and sedges from interdune meadows, such as
Panicum spp., Cenchrus incertus, and Cyperaceae, produce primarily dumbbell, small cross, and Cyperaceae
phytoliths. The Pooideae and Bambusoideae subfamilies
of grasses, which are generally common along the edges
or on the understory of maritime forests, produce wavy
trapezoid and long saddle phytolith, respectively. The
dominant grasses of swamps produce primarily dumbbell and plateau saddle phytoliths. Rondel/saddle
ellipsoid phytoliths are almost exclusively produced by
Spartina alterniora, the dominant plant in salt marshes.
Thus the predominance of rondel/saddle ellipsoid
phytoliths in a sediment sample may be a good indicator
of salt marsh environment. Rondel and spool/horned
tower phytolith are common in grasses from brackish
marshes.
This study provides a basis for the interpretation of
fossil grass phytolith assemblages recovered from
coastal sediments for the reconstruction of coastal
environmental changes. Some coastal environments,
such as salt marshes and brackish marshes, may not be
easily distinguishable based on their pollen assemblages
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