Michael Therin
11 Salisbury Road, Willoughby, NSW 2068, Australia
Robin Torrence
Division of Anthropology, Australian Museum, 6 College Street, Sydney NSW 2010, Australia
The analysis of starch grains preserved in sediments as a technique for palaeoenvironmental reconstruction is an
important new technique, but the limits of inference using this methodology still need to be assessed. A morphological
classification of starches extracted from topsoils collected from a range of modern environments in the humid tropical
region of Papua New Guinea was therefore conducted to see if discrimination among plots with dierent vegetation
could be achieved. Using an assemblage-based approach, starch grains, initially classified into morphological categories
and re-organized into groups according to size/shape frequency distributions, were grouped using principal components
analysis. The results correctly identified dierent types of human land use and provided good discrimination between
gardens and forest. These preliminary results show that past human land use can be satisfactorily reconstructed using
multivariate analysis of starch grain assemblages with only data on size and morphology to characterize the grains. The
findings of the starch study compared favourably with a phytolith analysis of material from the same locations,
although the two microfossils target dierent plant groups. Whereas phytoliths discriminate among dierent levels of
disturbance, starch grains can add information about specific site function, an important consideration especially for
regions where plant staples are typically high-starch producers. Combining starch grains and phytoliths in future studies
will enhance the accuracy of palaeoenvironmental reconstructions. 2002 Elsevier Science Ltd. All rights reserved.
PNG
0 100
km
Location
Map Bismark Sea
New Britain
Yombon
Kandrian Solomon Sea
Papua New Guinea Kumbum
Figure 1. Locations of soil samples used in the analyses (cf. Table 1).
in the composition of the starch assemblages Site Description and Sampling Procedures
from dierent layers, they proposed changes in local
vegetation and used these to infer site function. Their Sediment samples for the starch analysis were collected
conclusions were supported by additional archaeologi- from Garua, Kaula and Garala Islands and the Garu
cal data. Since the links made by Therin, Fullagar & village environs in West New Britain province, Papua
Torrence (1999) between starch grain size and vegeta- New Guinea. Fifteen localities, representing a variety
tion types were rather simplistic and in some cases may of habitats including ocean foreshore, closed forest,
have been incorrect, the main value of their study was regrowth forest, new gardens, fallow gardens, coconut
to demonstrate that starch assemblages can vary in plantations, and a village were sampled (Figure 1;
relation to changes in human land use. Further studies Table 1). At each location pinch samples of soil were
are required to evaluate how accurately and in what collected randomly within a 1010 metre quadrat.
ways the composition of starch grain assemblages in Soil from the uppermost 4 cm was included and litter
sediments monitors the plant communities from which was excluded.
the starch was derived. Local informants provided information about the
This article reports the results of a modern test case history of recent land use at each sampling locality,
which was carried out to measure the nature of the and a plant survey was carried out. Nine localities,
starch grain assemblages extracted from sediments Gu7ac, IG, Gu8, Gu3, Gu5, Gu11 and GMH3 had a
collected from plots with a variety of vegetation types cultivation history alternating between gardening and
and land use histories. Since cultivated food crops are bush-fallow. In the region traditional swidden tech-
characterized by large quantities of starch, we pre- niques are used for garden preparation. Shade trees,
dicted that, at the very least, variation in starch grain usually useful trees (e.g., mango, Canarium sp., bread-
assemblages should be able to dierentiate between fruits and figs) are usually left to grow and all other
cultivated plots and other types of vegetation vegetation is cut, piled into heaps and burnt. Taro
particularly forests in which starch-bearing plants are (Colocasia esculenta (L.) Schott) is usually the first crop
relatively dispersed and uncommon. planted on newly prepared beds, and following the first
In the absence of an adequate starch reference harvest other more resilient crops including sweet
collection, an assemblage-based approach of general potato, Singapore taro (Xanthosoma sagittifolium (L.)
morphologies using multivariate techniques was em- Schott), yams, bananas, cassava and sugar cane are
ployed. Previous studies have shown that this method- planted and allowed to grow with minimal follow-on
ology can process large amounts of data and establish care. Eventually garden plots are overtaken with
meaningful patterns of associations between micro- grasses, sedges, ferns and herbaceous weeds which give
fossils that show environmental dierentiation way to shade tolerant gingers, heliconias and ferns as a
(Powers, Padmore & Gilbertson, 1989; Boyd, Lentfer suite of colonizing trees (e.g. Mallotus, Ficus, Abroma,
& Torrence, 1998). Macaranga, Leukosyke, Endospermum, Omalanthus
Starch Grains and Environmental Reconstruction 689
Table 1. Summary of localities used for the starch and phytolith analyses. Localities marked by * were used for the
phytolith study. IG** was used for the starch analysis only
and Homalium species) create canopy cover. Plots are islands which were sampled. At these localities, K2 on
usually left to bush-fallow for 1020 years. Kaula Island, and G12 and G13 on Garala Island
Of the nine localities representing garden sites, four comprised palm forest with a mixed canopy of Euodia,
were freshly cleared areas planted with taro (Gu7c Myristica, Homalium and Diospyros species, an under-
and IG), one was an older garden with a ground cover storey dominated by palms, Caryota rumphiana Mart.
dominated by sweet potato vine (Gu5), and another and Hydriastele sp., and some small trees, mostly
(Gu3) was an abandoned garden with sugar cane, Polyscias cumingiana (Presl.) F.-Vill. Vines, including
bananas and cassava overgrown with a common Aster- Calamus ralumensis Warb., Capparis zippeliania Miq.,
aceae weed Ageratum conyzoides L. The remaining Smilax vitiensis A. DC., Faradaya sp. and Flagellaria
three comprised an abandoned garden site on Garua sp. were also common. The ground layer on the Garala
Island with well developed bush-fallow where gingers Island sites, G12 and G13 comprised mostly ferns,
dominated the ground layer and a tree common in gingers and heliconias. Palm seedlings were more
regrowth, Homalium foetidum (Roxb.) Benth. domi- common on the Kaula Island. In contrast to these
nated the canopy (GMH3), an area of forest that had localities, Nave River (Nv) comprised an inland forest
been freshly cleared after 10 years fallow where the recovering after logging approximately 12 years pre-
vegetation had been piled into heaps ready for burning viously. Common emergent and canopy trees included
(Gu8), and finally, another area immediately adjacent Euphorianthus longifolius Radk., Celtis sp., Myristica
to a swept village with grasses and coconut palms schleinitzia Engl., Chisocheton sp. and Pometia pinnata
(Gu11). J. R. and G. Ernst. The understorey comprised smaller
The other sampled localities did not have a recent trees and shrubs common in regrowth (Trema, Heriti-
garden history, although the two foreshore locations era, Pipterus species) and palms, mostly Hydriastele sp.
on Garua Island, FEK and GBL were within a coconut Gingers, ferns and aroids were common in the ground
plantation area characterized by ground cover consist- layer.
ing of grasses, ferns and herbaceous weeds. Local oral The variation in vegetation among the sampled
history states that minimal garden activity had oc- localities provides a good test case for whether starch
curred at some stage in the past on the small volcanic grains deposited in the soil can accurately reconstruct
690 C. Lentfer et al.
Table 2. Protocol developed by Therin (1998b) for extracting starch grains from sediments
III. Mounting
26. Add 500 l of ultra pure H2O to the dried suspension.
27. Vortex, using a pipette place 60 l on a microscope slide and let dry.
28. Mount in Euparol.
the local environment. The region has another advan- sodium polytungstate. The residue samples containing
tage for a test case of this type because the modern starch were mounted on to microscope slides in
topsoil is forming on a volcanic tephra that is known to Euparol. An Olympus BC60 light microscope with a
be no older than 1000 years (Machida et al., 1996). The polarizer was used to undertake the classification and
young age of the soil means that there is less chance of recording of the starch grains.
mixing from the various types of land use that might Using previous work undertaken by Therin (1994;
have occurred in the distant past (e.g. Torrence, Therin, Fullagar & Torrence, 1999), a key of common
Pavlides, Jackson & Webb, 2000). starch morphologies was drawn up (Therin, 1998c).
Individual starch types were defined using the follow-
ing features: hilium (visible/non-visible); position of the
Starch Extraction, Classification and hilium; lamellae (present/absent); two-dimensional
shape of the grain; and character and position of the
Counting Procedures extinction cross. Each combination of features was
Starch extraction from the sediment samples followed assigned an individual key number. As the analysis
a heavy liquid separation protocol (Table 2) developed proceeded, new types were added when encountered.
by Therin (1988b) on the basis of extensive experimen- In all, 90 separate starch types were included in the key
tation. This methodology represents a considerable (Figure 2).
improvement on the techniques used previously (e.g. The slides were initially scanned under polarized
Fullagar, Loy & Cox, 1998; Atchison & Fullagar, light at a magnification of 200 to locate the starch
1998; Therin, Fullagar & Torrence, 1999; Therin, grains for measurement. To ensure that duplicate re-
1994). The soil samples were oven dried at 40C and cording of starch grains was avoided, the slides were
five gram subsamples were taken for the analysis. scanned in transects the width of the field of view.
These were oxidized in hydrogen peroxide, sieved Recording of 100150 grains per sample was under-
through a 250 m mesh, deflocculated using Calgon taken at a magnification of 500. Both polarized and
solution, and fractionated for starch separation with non-polarized light were used as appropriate. The
Starch Grains and Environmental Reconstruction 691
1. 2.
81. Light yellow 84.
4. 6. 86. 87.
11. 12.
92. 97.
15. 16.
98. 99.
22. 23.
118. 119.
26. 27.
122. 123.
36. 38.
136. 137.
41. 44.
140. 141.
45. 46.
142. Light yellow 143. Light yellow
47. 48.
144. 145.
49. 51.
146. 147.
53. 55.
148. 149.
57. 59.
150. 151.
77. 79.
Figure 2. Expendable key of starch grain morphologies (Therin, 1998c). Grains are not drawn to scale.
shape and distinctive features of grains were noted according to morphological type using a number allo-
under non-polarized light. The character and location cated from the exandable starch key (Figure 2). Maxi-
of the extinction cross were then recorded under polar- mum length and width were recorded, with the width
ized light. Each starch grain was drawn and classified measured at the widest point perpendicular to the length.
692 C. Lentfer et al.
3
(a) Logged forest Small island sites
with palm forest
Nv
2 Garden, GI2
village and old
garden regrowth
sites
1
Gu7c Gu11
Coconut plantation Gu8
IG GI3
sites and garden
PC2
GBL
Gu3 GMH3
1
K2
2
3
2.5 2 1.5 1 0.5 0 0.5 1 1.5 2 2.5
PC1
0.6
1c 26c
(b)
57a
38a 139
0.4 47ab 15ab
26b 51 142
9bcd
79a 4a 12a
26a 27 18a 41a 99a
0.0 9a 1f 12bc 23c 4bc
36 16a 144a
6a 2de 92ab 60 128
98a 23ab 135a
76a
55
1a 49
0.2 22bc
2bc
28a
22a
89a
2a
0.6
0.8 0.6 0.4 0.2 0.0 0.2 0.4 0.6 0.8
PC1v1v2
Figure 4. Biplot of the first two principal components of the starch analysis explaining 33% of variation in the assemblages. A is the sample
plot and B is the vector plot. palm forest; old garden; new garden; coconut plantation;
regrowth forest; + village; logged
forest.
the variance. An examination of two bi-plots using left; and finally (4) gardens (IG, Gu3, Gu7ac), village
dierent combinations of these principal components (Gu11) and regrowth forest after gardening (GMH3
(PC1 versus PC2 and PC1 versus PC3, see Figures 4 & and Gu8), which are clustered in the middle of the
5) shows similar associations between sample localities. plots. It is notable that the coconut plantation localities
Four distinct habitat groups have been segregated by GFEK and GBL were from foreshore locations and
the analysis: (1) the logged forest (Nv) on the upper left although ground cover at these locations were domi-
of both plots; (2) the small volcanic islands with palm nated by ferns, grasses and other herbaceous weeds, it
forest (K2, G13 and G12) on the right of both plots; (3) is possible that at some time in recent history, Ipomoea
the coconut plantation plots (GFEK and GBL) and pes-caprae L. was present. This species typically forms
one of the gardens (Gu5) on the middle left and lower dense mats of ground coverage along foreshore sites in
694 C. Lentfer et al.
3
(a) Logged forest
Small island sites
Nv with palm forest
2
K2
Garden,
village and old
1 Coconut plantation garden regrowth
sites and garden sites
with Ipomoea
PC3
Gu8
Gu7c IG
0 GFEK Gu7a
GMH3
Gu3 GI3
Gu7b Gu11
GBL
1
GI2
Gu5
2
1.5 1 0.5 0 0.5 1 1.5 2 2.5
PC1
0.6
35
(b) 45a
33ab
47ab
0.4 26b
32a
1c 4bc
38b 40
145a
2bc
22a 22
89a
0.2 41a
11a
PC3v1v3
4a
12bc 128 135
1b 28a 26c 60
1a 1f
2a 4a 55 9bcd
36 76a 51
38a
0 79a 2de 144a 10a
6a 1d
23ab 23c
12a
20 77 92ab 15ab 57a 1e
16a 99a
138
81a 47c
137 143a
0.2 98a 53a 142
27a 139
9a
26a
18a
0.4
0.6 0.4 0.2 0 0.2 0.4 0.6 0.8 1
PC1v1v3
Figure 5. Biplot of the first and third principal components of the starch analysis explaining 31% of variation in the assemblages. A is the
sample plot and B is the vector plot. palm forest; old garden; new garden; coconut plantation;
regrowth forest; + village;
logged forest.
the region. It is a close relative of the sweet potato. types of vegetation and especially between places which
Therefore, this may explain the association of these have experienced swidden gardening and forests with
sites with the garden site Gu5, the only locality minimal or no gardening. In this analysis the regrowth
sampled with a dominant coverage of sweet potato forests on the small, volcanic islands and the logged
(Table 1). forest sample have been successfully separated from
the cultivated sites.
The position of the cluster in the middle of the starch
Discussion sample principal components plots (Figures 4 & 5)
The results confirm the expectation that variation in is notable. These localities alternate between food
starch assemblages does discriminate between dierent crops and regrowth during the current swidden system.
Starch Grains and Environmental Reconstruction 695
Consequently, they have diverse plant compositions In the meantime, the assemblage-based approach
typical of dierent environments: i.e. cultivated crops, has helped to establish the types of information that
herbaceous regrowth and tropical forest species. This the starch grains can provide and the results point to
partially accounts for the relatively low level of vari- areas that would benefit from further research. Multi-
ation in the assemblage explained by the first three variate analyses of starch grain assemblages can be
principal components (i.e. 45%). Although these sam- very useful as a first stage in classifying soil samples
pling localities have a variety of vegetation growing on into vegetation types, even if these cannot yet be
them now, they share a common history in terms of identified. The clustering resulting from the multi-
starch plants. All were once gardens that had tubers variate analysis suggests that the inter-assemblage dif-
with high starch content (taros, yam, and sweet po- ferences were related to land use. For many archaeo-
tato). It is therefore not surprising that these swidden logical studies it will be useful simply to know that
locations have all been grouped together. dierent environments are represented in the material
At this stage the division of shapes into subtypes under study and this may help establish further re-
based on size distribution is not definitive. Some search priorities. The patterns that result might also be
species probably have overlapping dimensions of further compared with soils taken from modern set-
starch grains and will have been ill-defined within this tings or with additional archaeological or plant micro-
analysis. For instance, Type 1 was split into 6 subtypes fossil data that could assist interpretation. To better
1a (15 m), 1b (69 m), 1c (1013 m), 1d (14 understand the patterns of variation, further research
22 m), 1e (2330 m) and 1f (3539 m) and each was can also be focused on the morphologies that are
associated with dierent categories of land use: 1a and associated with and define particular clusters and em-
1b had the strongest association with cultivated loca- phasis can be placed on identifying these important
tions (gardens and coconut plantation); 1c with the types to particular taxa first, rather than tackling
logged forest site; 1d with the palm forests of the small identification of whole assemblages.
volcanic islands; 1e with both forest and garden sites;
and finally, 1f was not clearly defined. The association
of 1e with both forest and garden sites may be due to Parallel study of phytoliths
incorrect splitting. Likewise, the subdivision of 1a and The results of the multivariate analysis of the starch
1b is questionable since 1a carries little weight within assemblages compare favourably with the previous
the analysis for the first 3 principal components (as phytolith analysis reported in Boyd, Lentfer &
indicated by its central position in vector plots) and it Torrence (1998), which was conducted in parallel to
is possible that the size range of this subtype was the starch grain research. Phytoliths were extracted
incorrectly determined. Alternatively, type 1a, being from sediments taken from many of the same localities
small and round and possibly common to many plant chosen for the starch grain study, but 14 additional
species, may occur in many environments and there- samples were included in the phytolith analyses (Table
fore cannot be used eectively to dierentiate between 1). Figure 6 represents a summary plot of the principal
them. Further statistical tests (e.g., Anova and Hom- components analysis of the phytolith data. In this case
ogeneity tests) could be applied to the size/shape dis- discrimination was mainly achieved by the respective
tribution to better define types and refine the overall distributions of grass, palm and arboreal phytolith
analysis and interpretation, but this preliminary analy- types (Boyd, Lentfer & Torrence, 1998). These types
sis has successfully demonstrated that starch grain separated sampling localities into groups representing
assemblages are capable of discriminating among dierent levels of clearing, cultivation and length of
various types of land use. fallow. For example, in Figure 6 the small island
regrowth forest sites with low levels of disturbance are
Assemblage based approach on the lower left of the plot; coconut plantations and
Since this analysis was conducted in the absence of a villages with high levels of disturbance and minimal
starch reference collection, it will certainly be much regrowth are on the right; disturbed locations associ-
improved when starch types can be assigned to plants. ated with large clearings and substantial regrowth
It has already been determined that many important are plotted in the middle of the plot; and disturbed
economic plants have diagnostic starch grains (e.g. localities associated with small forest clearings form a
Reichert, 1913; Loy, Spriggs & Wickler, 1992; Loy, loose cluster above middle of the plot.
1994; Piperno & Holst, 1998; Ugent et al., 1981, 1982, Dierences between the results of the phytolith and
1984, 1986, 1987) but very little work has been con- starch analyses are explained to a large degree by the
ducted on the thousands of species present in the study dierent plant groups targeted by each plant micro-
area. A productive approach in the long run is to begin fossil. Phytoliths are abundant in plants which are
the process of identifying the starch grains to plant prevalent in disturbed plant associations. For this
taxa through the examination of an appropriate refer- reason, the phytolith assemblages could not discrimi-
ence collection. This process has already been initiated nate eectively between the logged forest site, garden
for the West New Britain region (Therin, Torrence & sites, village sites, and sites dominated by herbaceous
Fullagar, 1997). regrowth, since grass presence was the major common
696 C. Lentfer et al.
3
(a) Disturbed sites
Gu7c associated with
small forest
clearings and
2 long forest fallow
Gu8
Gu6 Gu5
Gu9
1 Disturbed sites Gu7a
associated with large Nv
PC2
0.8
(b)
0.6 cys
bul3
long sm
0.4
misc
zing
0.2 ss
PC2v1v2
musaceae
block lpr
bul1
tracheid fac msps cyp tower
0.0 hb
ns bul2 fisp trap
ds
saddle trilob
0.2 lsps
bilob
cross
0.4
ssps
factor in these plant associations. In contrast, high The parallel analyses show that phytoliths are very
starch content is more typical of the food crops planted eective at determining levels of disturbance (Figure 6)
in the gardens and starchy plants are likely to be in terms of the extent of clearing and the nature
relatively rare in forests regardless of the amount of of cultivation (i.e., length of fallow in association
disturbance. The starch grain analysis was therefore with gardening and patterns of regrowth), whereas
successful at discriminating the logged forest and the starch analysis can confirm site function (i.e. garden
disturbed island forests, but the active and fallow versus non-garden). As a result, combining the two
gardens were all clumped together. studies will clearly enhance the reconstruction of
Starch Grains and Environmental Reconstruction 697
palaeoenvironments and human land use. This is National Museum and Art Gallery (PNG); West New
particularly relevant for contexts such as West Britain Provincial Cultural Centre and especially
New Britain, where the staple crops are high starch John Namuno; Kimbe Bay Shipping Agency; Garua
producers and mainly non-phytolith producers. Plantation; Walindi Plantation and Resort; Bill Boyd
and Ruth Henderson; and residents and land owners in
Garu Village, especially our guides Gabriel Loga and
Conclusions John Normu.
This study has shown that the analysis of starch grains
extracted from sediments can separate environments in References
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