People and plant interaction at the Houli Culture Yuezhuang site in Shandong Province, China

Abstract

Palaeoethnobotanical research at the Yuezhuang site, a Houli Culture settlement in Jinan, Shandong Province, China, dating to 8000–7700 cal. BP, documents human–environment interaction and the local subsistence economy soon after the initiation of food production in the region. This economy supported a sizeable community that occupied a kilometer stretch of floodplain along the Nandasha River. The research explores plant domestication, the extent to which the Yuezhuang population had developed a food production niche, and, to a lesser extent, the development of agriculture in the lower Yellow River valley. In order to do so, charred seeds from a variety of plant taxa were recovered by flotation of sediment from pits and cultural strata. Just over 30% of the seed assemblage is rice (Oryza sativa), broomcorn/common millet (Panicum miliaceum), and foxtail millet (Setaria italica subsp. italica). The status of several other plants such as soybean (Glycine max subsp. max or G. max subsp. soja), perilla (Perilla sp.), and chenopod (Chenopodium sp.) that are also cultivated in East Asia is also assessed. Most of the plant taxa are from open, sunlit, and anthropogenic, disrupted habitats. Diverse grasses similar to those found at later sites indicate that the farming niche documented at the late Neolithic Longshan Culture in Shandong Province was being established by 8000–7700 cal. BP. The plant remains assemblage is compared with three assemblages, two belonging to the Houli Culture and one from the late Neolithic Longshan Culture. Anthropogenic habitats and their formation, maintenance and use, wetland exploitation, cultivation, hunting–gathering–fishing, and animal management characterize the mature (late) Houli Culture niche.

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The Holocene
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DOI: 10.1177/0959683616650269
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Introduction
Yuezhuang is one of about 16 early-Holocene Houli Culture
occupations in Shandong Province, China (Figure 1) (Luan,
2009). Rescue excavations at Yuezhuang in 2003 provided a
valuable opportunity to explore human–plant interaction during
the early-Holocene when the lower Yellow River valley was rap-
idly evolving. Diverse subsistence strategies were emerging in
Northeast Asia by 8000 years ago, when Yuezhuang was occu-
pied. These subsistence strategies ranged from apparent hunting
and gathering (e.g. early Chulmun of the Korean Peninsula, Ini-
tial Jomon in Japan), to mixed economies that emphasized hunt-
ing and gathering and resource management. In China, arboreal
perennials through annual plants were likely managed as were
some early crops such as millets, rice, peach, and probably soy-
bean (Cohen, 2011; Crawford, 2011; Lee, 2011; Zhao, 2011;
Zheng et al., 2014). Preliminary results of the Yuezhuang flota-
tion sample analysis have already added to the growing knowl-
edge of early-Holocene subsistence regimes in North Asia
(Crawford et al., 2006, 2013), placing it well along the contin-
uum between hunting–gathering and farming. Yuezhuang has
been notable because it has the earliest rice (c. 8000 cal. BP) so
far reported in the Yellow River basin (Crawford et al., 2006).
The charred rice grains may be evidence of its local cultivation in
the region, exchange from the south, or that it was a wild resource
(see Jin et al., 2014 for a brief discussion). We have always con-
sidered the implications of the rice to be ambiguous and its role
at Yuezhuang in need of further testing (Crawford et al., 2013).
Although previous summaries of the palaeoethnobotanical
research at Yuezhuang reported the rice and other plant taxa, the
full implications of the collection have not been explored nor has
the complete dataset been reported.
Two other archaeobotanical assemblages have recently been
reported from sites closely related to Yuezhuang, Zhangmatun,
and Xihe sites (Jin et al., 2014; Wu et al., 2014). All three sites
include a wide range of plant taxa among the charred remains
People and plant interaction at the
Houli Culture Yuezhuang site in
Shandong Province, China
Gary W Crawford,1 Xuexiang Chen,2 Fengshi Luan2
and Jianhua Wang3
Abstract
Palaeoethnobotanical research at the Yuezhuang site, a Houli Culture settlement in Jinan, Shandong Province, China, dating to 8000–7700 cal. BP,
documents human–environment interaction and the local subsistence economy soon after the initiation of food production in the region. This economy
supported a sizeable community that occupied a kilometer stretch of floodplain along the Nandasha River. The research explores plant domestication,
the extent to which the Yuezhuang population had developed a food production niche, and, to a lesser extent, the development of agriculture in the
lower Yellow River valley. In order to do so, charred seeds from a variety of plant taxa were recovered by flotation of sediment from pits and cultural
strata. Just over 30% of the seed assemblage is rice (Oryza sativa), broomcorn/common millet (Panicum miliaceum), and foxtail millet (Setaria italica subsp.
italica). The status of several other plants such as soybean (Glycine max subsp. max or G. max subsp. soja), perilla (Perilla sp.), and chenopod (Chenopodium
sp.) that are also cultivated in East Asia is also assessed. Most of the plant taxa are from open, sunlit, and anthropogenic, disrupted habitats. Diverse
grasses similar to those found at later sites indicate that the farming niche documented at the late Neolithic Longshan Culture in Shandong Province
was being established by 8000–7700 cal. BP. The plant remains assemblage is compared with three assemblages, two belonging to the Houli Culture
and one from the late Neolithic Longshan Culture. Anthropogenic habitats and their formation, maintenance and use, wetland exploitation, cultivation,
hunting–gathering–fishing, and animal management characterize the mature (late) Houli Culture niche.
Keywords
anthropogenesis, archaeobotany, China, early-Holocene, early Neolithic, flotation, Houli Culture, millet, niche construction, origins of agriculture, rice,
Shandong Province
Received 3 February 2016; revised manuscript accepted 25 April 2016
1 Department of Anthropology, University of Toronto Mississauga,
Canada
2 Department of Archaeology and Museology, Shandong University,
China
3
School of Tourism, History and Culture, Southwest University for
Nationalities, China
Corresponding author:
Gary W Crawford, Department of Anthropology, University of
Toronto Mississauga, Mississauga, ON L5L 1C6, Canada.
Email: g.crawford@utoronto.ca
650269HOL0010.1177/0959683616650269The HoloceneCrawford et al.
research-article2016
Research paper
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2 The Holocene
(including crops) indicating at the very least a broadly based plant
exploitation strategy in the region between 9000 and 7700 cal. BP.
We document the full charred plant remains dataset from
Yuezhuang here in order to address two questions: (1) to what
extent are the two millet species and rice domesticated and (2)
where along the trajectory to food production does the Yuezhuang
subsistence economy sit? To answer the second question, we
present a model for Houli Culture plant and human interaction at
the onset of the Neolithic in the region that is similar in many
ways, but not identical, to that of the late Neolithic in Shandong
Province. Our results inform the discourse on human–environ-
ment interaction, anthropogenesis, and niche construction/eco-
logical engineering in North China.
The Yuezhuang site
Yuezhuang is a Houli Culture occupation situated 7.3 km west of
the Yellow River on the south side of one of its tributaries, the
Nandasha River, and sits 64 km southeast of Xihe, a site that is
contemporaneous with Yuezhuang, and 42 km southeast of the
9000–8500 cal. BP Zhangmatun site (Wu et al., 2014) (Figure 1).
A prolonged period of landscape stability at the Yuezhuang locale
began c. 8000 cal. BP following a 5000-year episode of alternat-
ing erosion and deposition that began to stabilize c. 9000 cal. BP
(Zhuang et al., 2013). Low mountains, part of the Shandong
Highlands, rise to just over 200 m about 3 km to the east. The
Houli Culture occupies an east–west corridor in Shandong Prov-
ince west of Changbai Shan, a mountain range, which stretches
from Jinan to Yantai (Luan and Wagner, 2009: 3).
The Houli Culture dates to c. 9000–7500 cal. BP (Luan and
Wagner, 2009: 4; Wu et al., 2014) and is the earliest Neolithic
presence in Shandong Province. Radiocarbon dates from the Xihe
and Yantai Glassworks sites range from about 10,000 to 7400 cal.
BP, but the earliest and latest dates are considered outliers. In all,
10 AMS dates from the early Houli Culture Zhangmatun site
range from c. 9000 to 8500 cal. BP (2σ) (Wu et al., 2014). Seven
AMS dates from Xihe on seeds and stem fragments range from
8025 to 7575 cal. BP (2σ) (Jin et al., 2014). Two direct AMS
radiocarbon dates on rice and millet grains from Yuezhuang are
7050 ± 80 BP (TO-11865) and 6900 ± 35 BP or c. 8000–7700 cal.
BP (BA-8168), respectively (2σ) (Figure 2). Rice is also reported
from Xihe; four grains are AMS-dated to 8025–7850 cal. BP (Jin
et al., 2014).
Broomcorn and foxtail millets are not unique to Yuezhuang,
having been recovered from other Houli Culture sites (Jin et al.,
2014; Wu et al., 2014). Soybean and perilla are also part of the
Yuezhuang assemblage indicating that a close association of peo-
ple with plants that would eventually become important crops in
East Asia was developing. Animal remains from Yuezhuang are
diverse and include fish, reptiles, small and large mammals
(mainly deer, cattle, and pig), and birds; among these, the dog and
pig at Yuezhuang are considered domesticated (Song, 2003).
Figure 2. AMS radiocarbon date probability curves for rice and millet recovered from the Yuezhuang site. The thick and thin horizontal lines
indicate the 1σ and 2σ probabilities, respectively. The dates were calibrated using Calib 7.1 (Stuiver and Reimer, 1993).
Figure 1. Regional map showing archaeological sites mentioned in
the text.
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Crawford et al. 3
Several periods of occupation are recognized at Yuezhuang, but
the samples reported here are mainly from the earliest period.
Strata 6 and 7 are Eastern Zhou cultural layers (Center for East
Asian Archaeological Studies, Shandong University (CEAASSU),
2005). Four areas were excavated; three are located in or near the
northernmost area that was disturbed by a brick factory. The fourth
area (T0988 to T1186, T1085) is about 800 m to the south of the
three other excavation units (CEAASSU, 2005). The excavations
exposed numerous pits, none of which appear to be associated with
dwellings. Pits associated with ditches (G3 and G4) were uncov-
ered only in the southern unit (Figure 3). The pottery is mostly
undecorated and sand tempered. Ground and polished axes and
adzes, grinding stone (querns), bone points, awls, and unknown
tools are all typical of the early Neolithic in the region.
Methods
Sediment from 35 pits and several non-pit strata resulted in flotation
samples from 55 contexts (Tables 1 and 2; Figure 3). One sample is
from a Zhou period context and several others are from two layers
that appear to be natural deposits or strata that disclosed no clear
occupation evidence. None of the non-Houli samples contain more
than four charred seeds. The samples were processed using the light-
weight, rectangular device that we call the ‘modified SMAP (Shell
Mounds Archaeological Project) device’ (D’Andrea et al., 1995).
Light and heavy fractions were stored in cloth and dried. The result-
ing material was not abundant, so most samples were not screened
before being examined, as would normally be the case (Crawford,
1983). The light and heavy fractions were examined at Shandong
University using a Nikon SMZ1000 microscope. The specimens
were photographed with a Nikon Digital Sight camera system (DS-
5M-L1). Larger specimens were weighed and counted; otherwise
smaller seeds and other fragments were only counted.
Results
The flotation sampling recovered 453 small seeds and a few frag-
ments of both acorn meat (cotyledon) and shell (pericarp or seed
Figure 3. Contexts sampled at the Yuezhuang site with seed densities indicated by the bubble diameters. Bottom right is the southernmost
excavation unit with ditches.
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4 The Holocene
Table 1. Numbers and weights of seeds by taxa and context recovered by flotation from the Yuezhuang site.
Sample Weights (g) Crops Weedy grasses Other weedy annuals Fleshy
fruit
Other
seeds
Totals
Volume
(liters)
T
(Unit)
H
(Pit)
Layer
Total wt (g)
Unidentifiable
remains
Wood
charcoal
Acorn meat
Acorn shell
Seeds
Rice
Foxtail millet
Broomcorn
millet
? millet
Green foxtail
grass type
Paniceae
Paniceae 1
Panicum sp.
type
Barnyard grass
Brassicaceae
Chenopod
Knotweed
Pale
smartweed
Legume
Soybean
Portulaca
Perlla
Grape
Unidentifiabe
Unknown
Total seeds
Total seed
density (no./L)
36 6051, 6052 61 * * 3 1 2 3 10 1 11 1 32 0.9
6 6153 72 * 2 2 0.3
12 5443, 5445 80 1 1 2 0.2
12 5344 81 9 1 4 21 6 41 3.4
6 6053 90 1 1 1 3 0.5
6 6153 91 * * 0 0.0
12 6052 92 4 1 3 1 1 2 12 1.0
6 1186 123 1 1 2 0.3
6 6153 124 0.40 0.01 0.06 0.2 0.13 26 2 1 2 1 32 5.3
6 5344 126 1 1 0.2
6 5344 127 3 1 4 0.7
12 5344, 5444 129 1 1 1 2 5 0.4
6 5344 135 6 6 1.0
12 5344 136 6 5 1 6 2 20 1.7
12 5344 137 1 2 1 4 0.3
18 5344 138 8 6 3 7 24 1.3
6 5344 145 2 2 0.3
36 5344 146 2 33 3 7 4 2 4 1 1 1 26 4 88 2.4
6 1186 148 1 7 2 10 1.7
6 5344 151 1 1 0.2
6 6051 152 * 1 1 8 1 11 1.8
6 5344 154 * 0 0.0
6 5344 156 1 1 0.2
6 5344 157 1 4 1 3 9 1.5
6 5344 158 1 1 1 9 1 13 2.2
6 5344 159 1 1 1 30 3 36 6.0
6 6152 162 * 1 1 0.2
6 5934 167 1 1 0.2
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Crawford et al. 5
Sample Weights (g) Crops Weedy grasses Other weedy annuals Fleshy
fruit
Other
seeds
Totals
Volume
(liters)
T
(Unit)
H
(Pit)
Layer
Total wt (g)
Unidentifiable
remains
Wood
charcoal
Acorn meat
Acorn shell
Seeds
Rice
Foxtail millet
Broomcorn
millet
? millet
Green foxtail
grass type
Paniceae
Paniceae 1
Panicum sp.
type
Barnyard grass
Brassicaceae
Chenopod
Knotweed
Pale
smartweed
Legume
Soybean
Portulaca
Perlla
Grape
Unidentifiabe
Unknown
Total seeds
Total seed
density (no./L)
12 6034 172 * 3 1 1 2 1 4 2 14 1.2
12 6034 179 * 1 2 1 1 1 6 0.5
6 5934 182 1 2 1 1 7 12 2.0
18 5344 189 15.24 0.03 15.21 * 1 1 1 3 3 3 1 13 0.7
6 5344 191 1 1 0.2
6 6034 195 1 1 2 0.3
6 6151 225 1 1 3 5 0.8
6 5344 6 1 1 1 2 1 4 0.7
6 6153 6 1 1 0.2
6 5344 7 2 2 0.3
12 6052 7 4 1 5 0.4
12 5344 8 1 1 2 0.2
6 5344 9 3 3 0.5
30 6053 9 1 2 3 0.1
12 5344 10 * 1 1 2 0.2
6 6052 10 1 1 0.2
6 6150 10 * 1 1 0.2
6 6051 11 1 2 1 4 0.7
6 6052 11 1 1 0.2
6 6053 11 1 1 2 0.3
6 6153 11 1 1 0.2
6 6051 * 1 1 1 3 0.5
12 6152 1 1 2 0.2
Total 486 15.74 0.04 15.27 0.10 0.20 0.13 28 9 91 13 1 28 11 2 2 1 41 3 1 3 7 1 5 1 174 33 453 0.9
*Less than 0.01 g.
Table 1. (Continued)
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6 The Holocene
coat) (Figure 5). The overall seed density at the site is about 1
seed per liter. The samples with the highest densities are from
H146 (5.3 per liter) and H159 (6.0 per liter) (Figure 3). The plant
remains comprise 18 plant taxa that have been identified to fam-
ily, tribe, genus, species, or subspecies (Table 2). The actual num-
ber of taxa is probably more than triple this if we include the as
yet unidentified unknown specimens (9% of the assemblage
appears to be identifiable but as yet are not known); we estimate
one taxon per unknown of which there are 33 in addition to
unidentified grasses. About 37% of the specimens are unidentifi-
able because of breakage, charring, and/or lack of diagnostic fea-
tures. This is not unusual in a small collection like this one. Many
taphonomic influences result in the preservation of charred seeds
in archaeological deposits, and the process of creating this record
is far from perfect.
Crops, notably rice, broomcorn (common) millet, foxtail mil-
let, or unidentifiable millet, comprise 31% of the seeds (Figure 4).
Unidentifiable millet grains are so fragmentary that the embryo
shape and overall form of the original grain are difficult to assess.
Rice has been recovered from only two contexts. All but 2 of the
28 grains are from H124 where they are about 81% of the seeds in
H124. No rice spikelet bases have been recovered, so we cannot
assess the degree of shattering, a trait that is a clear indicator of
selection and management. Broomcorn millet, soybean, and
perilla comprise the other plant taxa identified in H124. The other
two rice grains are from H146 where 97% of the 77 seeds are mil-
let, wild grass, and a legume. None of these associations appears
to be significant.
Five other types of grasses, all in the Paniceae tribe, comprise
10% of the seeds (Figure 4). Many can only be identified to the
Paniceae tribe because the tribe is represented by a large number
of genera in China with considerable morphological overlap.
Some resemble Panicum spp., so they are listed as Panicum type.
Panicum has 21 species in China, among which 4 are introduced
(Chen and Renvoize, 2006). Among the endemic species, all are
listed as sub-tropical to tropical except Panicum bisulcatum.
Yuezhuang was occupied at the beginning of the hypsithermal, so
to assume that only P. bisulcatum was growing in the area at the
time is probably not wise. These Panicum-type seeds do not
resemble P. bisulcatum.
Other charred Paniceae caryopses in the samples are similar to
green foxtail grass, the wild ancestor of foxtail millet. Two barn-
yard grass seeds are in the sample from H146. This taxon is quite
rare in flotation samples in China. They are quite common in flo-
tation samples from Japan where we have evidence of its domes-
tication (Crawford, 1983, 1992). Paniceae Type 1 is represented
by 11 specimens in four pits and one cultural layer sample. It is
rectangular in dorsal view with an embryo that is less than half the
length of the seed. The seed surface has a distinctive sculpting
(Figure 4).
Non-grass, weedy, herbaceous annual plants represent about
13% of the assemblage: chenopod, knotweed, pale smartweed,
mustard family, perilla, and purslane. Chenopodium is repre-
sented by two seed types: one is a smaller form with a thick seed
coat (testa) and the other is larger with a thin seed coat (Figure 4).
A total of 14 species are reportedly endemic to China (Wu and
Raven, 2003). Species that appear to be distributed in north tem-
perate China include C. glauca, C. acuminatum, C. gracilispicum,
C. urbicum, C. ficifolium (a possible candidate for the small che-
nopod seeds), C. strictum, and C. album. C. giganteum is widely
cultivated in China and is considered part of the C. album aggre-
gate (Wu and Raven, 2003). Problems with the taxonomic history
of this genus make assessment of its status and identification dif-
ficult. Modern seeds have a reticulate-lineate sculpting and are
about 1.5 mm in diameter. The larger chenopod seed type at
Yuezhuang is smaller than this but has reticulate-lineate sculpting
Figure 4. Charred plant remains recovered from Yuezhuang: (a)
Perilla sp.; (b) two types of Chenopodium sp.; (c) Polygonum/Persicaria
lapathifolium; (d) foxtail millet; (e) broomcorn (common) millet; (f)
soybean; (g) grape; (h) small Paniceae grasses (1a and b: Echinochloa
crus-galli; 2: probable Digitaria; 3–7: rectangular grass with short
embryos; 8: possible Setaria italica subsp. viridis type; 9: Panicum-type
sp.; 10: unidentifiable grass); (i) acorn meat (cotyledon); and (j) rice:
lateral (left) and dorsal (right) views of rice seeds from pit H124. All
images are to scale except Perilla (a).
Table 2. Scientific and common names of plant taxa identified in
the Yuezhuang flotation samples.
Common name Scientific name
Cultigens
Broomcorn millet Panicum miliaceum
Foxtail millet Setaria italica subsp. italica
Rice Oryza sativa
Possible crops
Soybean Glycine max subsp. max/G. max subsp. soja
Beefsteak plant Perilla sp. ( P. frutescens or a diploid ancestor)
Chenopod, goosefoot Chenopodium sp.
Weedy plants
Bean family Fabaceae
Knotweeds Polygonum sp. and Persicaria sp.
Pale smartweed Polygonum lapathifolium
synonym: Persicaria latifolia
Mustard family? Brassicaceae
Purslane Portulaca sp.
Grass family Poaceae
Barnyard grass Echinochloa crus-galli
Foxtail grass type Setaria italica subsp. viridis type
Panic grass type Panicum sp. type
Millet tribe Paniceae
Fleshy fruit
Grape Vitis sp.
Tr e e
Acorn Fagaceae: Quercus/Cyclobalinopsis sp.
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Crawford et al. 7
(Figure 4). Archaeobotany may provide one way to explore the
history of chenopod cultivation in China. We are not claiming that
the Yuezhuang chenopod was cultivated, but similar seeds occur
at later Neolithic sites and its relative abundance at Yuezhuang
suggests that this possibility should be left open. Chenopods are
the second most abundant of the seeds in the assemblage, follow-
ing behind broomcorn millet. The seeds and greens of plants in
this genus are edible and nutritious. In the New World, several
species were domesticated and seed coats in domesticated forms
are quite thin, less than 10 µm (Smith, 1984, 1992). The seed coats
of the larger, reticulate chenopod seeds have not been measured,
but they are similar to ones recovered from Liangchengzhen. The
distribution of the chenopod and millet appears to be independent,
suggesting that they were deposited as a result of separate pro-
cesses. If two outlier samples, one with 8 chenopod and one with
18 broomcorn millet, are removed from the analysis, then the cor-
relation is stronger.
Four knotweed seeds (actually achenes) are represented by
three unknown species and one pale smartweed specimen (Figure
4), a common weed along roadsides, ditches, field margins, and
watersides. Knotweeds provide edible greens and small seeds. A
single possible mustard family seed is in one sample. The size and
shape is consistent with Brassica sp., but the seed coat pattern is
not at all clear and the specimen is fragmentary. Little significance
should be given to this seed. It may be easily confused with perilla.
Four specimens of Perilla sp. are in three samples (Figure 4).
Perilla is a crop with a long history of cultivation in Japan, Korea,
and China. Perilla has two varieties, an oil seed variety (P. frutes-
cens var. frutescens) and a vegetable (P. frutescens var. crispa).
The crops are tetraploid, one contributor being P. citriodora; two
other diploid species of perilla are recognized, but whether they
contributed to P. frutescens is not known (Nitta et al., 2003). The
Yuezhuang specimens are quite small, about 0.5 mm in diameter,
much smaller than either cultivar. We are not able to identify them
more specifically. Nevertheless, perilla is associated with the
anthropogenic habitats at Yuezhuang and we cannot rule out its
cultivation here.
A plant normally associated with extreme disturbance such as
cultivation is purslane. However, only one charred purslane seed
was recovered, so its significance at Yuezhuang is low. Several
species of purslane are considered endemic to tropical regions of
China, but only one is pervasive (in temperate and tropical
regions): Portulaca oleraceae. The species is well adapted to
open, disturbed habitats, particularly cultivated fields and the suc-
culent leaves are a nutritious vegetable.
At least two specimens are soybean (Figure 4). The hilum,
hypocotyl-radicle, shape, and symmetry of the beans are typical
of the genus. Seven other legume seeds have the general charac-
teristics of Glycine sp. but the hilum and radicle are not clearly
discernible. These are also listed in the soybean column in Table
1. The confirmed soybean specimens are smaller than later Neo-
lithic soybean (Lee et al., 2011); however, their size does not
help diagnose the domestication status of the soybeans because
of the size variation among wild, domesticated, and in-between
(including crosses) soybean. The characteristics that would bet-
ter distinguish wild from early domesticated soybean such as
growth habit and degree of pod dehiscence are archaeologically
invisible.
Discussion
The charred seed assemblage from Yuezhuang helps inform not
only what plants were available to, and potentially used by, peo-
ple during the Houli Period, but also the nature of human–plant
Figure 5. Box plots of length (L), width (W), thickness (T), length/width (L/W), and width/thickness (W/T) for broomcorn and foxtail millet.
The circles graphically represent the results of Student’s t-test. The non-intersecting circles for broomcorn millet (the centers of which are the
means of each population) indicate that the means are significantly different. The overlapping circles for foxtail millet indicate that the means
are not significantly different. The horizontal line is the grand mean, the box delineates the 25th and 75th percentiles, and horizontal lines
immediately outside the boxes represent the standard deviation.
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8 The Holocene
interactions at the time. The following discussion explores spe-
cific issues related to these interactions: the extent to which spe-
cific plants such as millet, rice, soybean, perilla, and chenopod
were undergoing selection for domestication syndrome traits, the
human–ecological context at Yuezhuang, and the extent to which
it involved food production, and how the assemblage compares to
other early Neolithic charred plant assemblages in North China.
The crops
Yuezhuang contains substantial remnants of two crops, broom-
corn millet and rice. Only 9% of the millet grains is foxtail millet.
The more numerous charred broomcorn millet grains are among
the oldest directly dated millet grains in China (Figure 2). Other
millet grains of a similar age are reported from sites such as Cis-
han, Dadiwan, Xinglonggou, Xihe, and Zhangmatun (Figure 1)
(Liu et al., 2004; Lu et al., 2009; Zhao, 2005). Only chaff is
reported from Cishan, so the millet grain morphology there is not
evident. We compare the Yuezhuang broomcorn millet measure-
ments to those of charred archaeological examples from the c.
1200–1300 cal. BP Satsumon period, Kashiwagigawa site in Hok-
kaido, Japan (Crawford, 1993), because they are the youngest
charred archaeological specimens in our collections and therefore
they best represent a very recent phenotype preserved under simi-
lar conditions. The Yuezhuang broomcorn millet grains (averag-
ing 1.5 mm long, 1.0 mm wide, and 1.0 mm thick), are significantly
shorter, narrower, and thinner (dorsally compressed) than the rela-
tively modern grains (Figure 5). A characteristic of broomcorn
millet is its spheroidal ventral and dorsal shape. Grains are also
relatively thick (not dorsally flattened). The box plots (Figure 5)
compare the shapes of the broomcorn millet populations. The
Kashiwagigawa millet grains are significantly more spheroidal
than the Yuezhuang grains. The grains are also shorter and nar-
rower than the specimens from Xinglonggou; however, the thick-
nesses of the Yuezhuang and Xinglonggou specimens are similar
(Figure 5) (Zhao, 2005). They are slightly smaller than the Zhang-
matun grains (Wu et al., 2014), but only six grains have been
recovered from Zhangmatun, so the size difference is probably
not significant. The lengths and thicknesses of the Yuezhuang
grains overlap with the recent, domesticated phenotype, so they
have undergone some selection. The ancestors of the tetraploid
domesticated broomcorn millet are still ambiguous (Hunt et al.,
2011, 2014), so we do not have reference seeds for a precise
ancestor with which to compare the Yuezhuang grains. Neverthe-
less, the thick grains combined with their quantity and distribu-
tion at Yuezhuang support the cultivation hypothesis. The
specimens from Yuezhuang appear to be an early form of the
crop, similar to the millet at Xinglonggou (Zhao, 2005). Selection
had not yet produced the larger, spheroidal phenotype typical of
recent broomcorn millet.
Foxtail millet grain metrics average 1.2 (L), 1.2 (W), and 0.9
(T) mm (Figure 5). The box plots (Figure 5) illustrate that the
Yuezhuang specimens fall within the range of variation of the later
phenotype, evidence that foxtail millet had already gone through a
significant degree of selection by c. 8000–7700 cal. BP. The
Zhangmatun foxtail millet measurements are problematic (averag-
ing 1.66 by 1.33 by 1.16 mm) (Wu et al., 2014) because they are
unusually long, being longer than most of the charred sample from
Japan. The photo of foxtail millet from Zhangmatun (Wu et al.,
2014), however, suggests that the grains are similar in size and
shape to the examples from Yuezhuang. Both the foxtail and
broomcorn millet grains from Yuezhuang and Zhangmatun are
within the morphological range of the crop forms indicating that
their domestication began some time before 9000–8000 cal. BP.
The rice at Yuezhuang was likely a form that was cultivated,
although where it was cultivated is not clear because we cannot be
certain that it was grown locally. The measureable grains from
Yuezhuang and Xihe are shorter than 5.5 mm and narrower than
2.7 mm (Crawford et al., 2013), a size that overlaps with wild and
domesticated phenotypes (Zhao and Gu, 2010). Where the Houli
Culture rice sits along the continuum from the wild to domesti-
cated forms cannot, therefore, be determined. Only a small por-
tion of the rice grains at Jiahu, a site that is partly contemporary
with Yuezhuang, are longer and wider than 5.5 and 2.7 mm,
respectively, so their domestication status is also difficult to
resolve (see Liu et al., 2007: 1065). This is not surprising consid-
ering that the age of these rice populations situates them early in
their domestication. Seed size changes result from complex influ-
ences that likely postdate selection for other traits such as non-
shattering (Fuller, 2007; Zhao, 2010b). The lack of rice spikelet
bases means that we cannot assess the degree of shattering, a trait
that is a clear indicator of selection and management. Neverthe-
less, both older and coeval rice remains from the lower Yangzi
valley indicate that selection for domestication related traits was
well under way.
The cluster of rice in one feature contrasts with the distribution
of millet and chenopod that are found in the majority of the pits,
occasionally occurring in clusters. This suggests a contrasting
cultural or subsistence value for rice. However, rice could easily
have been as important (materially and symbolically) as the other
crops. At the Longshan Culture Liangchengzhen site, for exam-
ple, a cluster of rice was found in one pit but is in low densities
elsewhere in many other contexts throughout the site (Crawford
et al., 2005). Furthermore, rice chaff in Liangchengzhen pottery is
evidence that rice had broader significance (including technol-
ogy) there than it did at Yuezhuang. However, until pottery manu-
facturing at Yuezhuang is examined, this comparison is tentative.
Yuezhuang and Xihe are located north of what appears to be the
natural range of the wild ancestor of domesticated rice. The closest
reports of archaeological rice contemporary with Yuezhuang and
Xihe are from Jiahu and Kuahuqiao, about 420 and 770 km away,
respectively. Archaeobotanists have argued that the rice at Kua-
huqiao was managed by, and undergoing domestication by, people
based on the fact that roughly half of the rachises are from a non-
shattering morphotype. The debate concerning whether the rice is
domesticated at Kuahuqiao has some impact on our interpretation
of the rice at Yuezhuang; human selection had significantly shifted
the balance of the rice population to one that is largely adapted to
human exploitation so that the rice at Yuezhuang should be simi-
larly adapted. Kuahuqiao is located well within the natural distri-
bution of wild rice, but the evidence points to Yuezhuang being
well north of the distribution of any species of wild rice. It was
difficult for people to isolate breeding populations of rice that they
were cultivating at Kuahuqiao (although they clearly had some
success), so crosses between wild rice (non-cultivated) and culti-
vated rice must have been common. Taking rice northward to sites
such as Yuezhuang may have been an important step toward an
unintended consequence: the isolation of managed or domesti-
cated rice from its wild ancestor.
Another consequence of bringing rice to the Yellow River val-
ley was to bring it into close association with foxtail millet. Rice
blast, a fungal parasite of rice, was originally a foxtail millet para-
site that shifted to rice when the opportunity was presented
(Couch et al., 2005). Yuezhuang is the earliest assemblage with
the presence of both foxtail millet and rice, so the shift potentially
occurred in the Houli Culture. Nevertheless, rice was likely an
important plant to the Yuezhuang people. It was either cultivated
or at the very least it was being exchanged into the region.
Three other plants that document a long association with peo-
ple in China are represented in the Yuezhuang samples: cheno-
pod, soybean, and perilla. Chenopod and soybean are reported
over an extensive geographic area and chronological span in
China. Perilla, although less common than chenopod and soybean
in the Chinese archaeological record, is still important because of
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Crawford et al. 9
its early association with people at Yuezhuang, the earliest report
of which we are aware. At the moment, whether initiation of their
cultivation had begun by 8000 years ago is an open question. For
the time being, we do not take a position on their domesticated
status because of the arbitrary nature of the definition (see Zeder,
2006 for a full discussion). This means that we do not make an a
priori assumption that they are wild either.
Early Neolithic plant use
The archaeological record for late 9th and early 8th millennia cal.
BP in north and central eastern China exemplifies adaptations to
interior riverine habitats and nearby uplands (Cishan, Dadiwan,
Peiligang), uplands consisting of rolling hills (Xinglonggou, Xin-
glongwa), slightly elevated plains surrounded by low mountains
(Shangshan, Xiaohuangshan), and low wetlands (Jiahu, Kuahuq-
iao). Yuezhuang, Cishan, Dadiwan, and Peiligang, while they are
interior, riverine associated sites, are on terraces at the edge of
deep loess deposits (see Chen, 2004). These locations facilitated
access to a rich array of animal and plant resources. All of these
cultures have evidence of food production. Five northern tier,
early Neolithic, sites have comprehensive flotation sample analy-
ses reported – Dadiwan, Xinglonggou, Yuezhuang, Zhangmatun,
and Xihe (Jin et al., 2014; Liu et al., 2004; Wu et al., 2014; Zhao,
2005) – while other analyses are ongoing. The Dadiwan plant
remains report is preliminary, focusing on the sequence of crops
in the five periods at the site. Flotation samples from Jiahu, on the
boundary between the northern and southern tier of sites, have
also been reported (Zhao, 2010a). Rice is present in the Jiahu
samples, but no millet was recovered (Zhao, 2010a: 106). Wild
grasses recovered from Jiahu include Echinochloa and Paniceae.
Yuezhuang and Jiahu also have chenopod, knotweed, soybean,
grape, and acorn. Otherwise, the Jiahu remains indicate that wet-
lands were significant plant foraging areas while we have no or
little evidence for wetland plants (other than rice) being important
to the Yuezhuang occupants. The occupants of Locality 1 at Xin-
glonggou grew primarily broomcorn millet but a few weeds (but
no tree fruit) are also in the samples (Cerastium, Astragalus, and
Chenopodium). Only broomcorn millet is reported in the earliest
period at Dadiwan while foxtail millet appears in later occupation
layers there, c. 6000 years ago (Liu et al., 2004). Similarly, broom-
corn millet is the first crop at Cishan, followed by foxtail millet
(Lu et al., 2009).
These sites all lie in the early part of a continuum from hunt-
ing–gathering to intensive agriculture. Although food production
at these sites was not as intensive as it was at middle and late
Neolithic sites, the evidence points to a significant commitment to
food production as part of a subsistence system that included
hunting, fishing, and gathering. Xinglonggou seems more focused
(i.e. a very narrow range of plants is represented) than either
Yuezhuang or Jiahu. The main qualitative difference between
Jiahu and Yuezhuang appears to be the importance of wetland
resources at Jiahu.
The Yuezhuang niche
People, like many other organisms, engineer their environs to
varying degrees for particular purposes and other organisms adapt
to, evolve, or cannot maintain populations in this context. This is
niche construction (Bleed, 2006; Crawford, 2014; Odling-Smee
et al., 2003; Smith, 2007). Furthermore, anthropogenic influences
range from directed to inadvertent (Crawford, 1997). Plants are
particularly sensitive indicators of these activities and coevolu-
tion in these contexts is best evidenced by domestication, a pro-
cess that clearly involves the Houli Culture. A conceptualization
of this region’s ecological engineering trajectory can be devel-
oped by comparing the assemblages from the three neighboring
Houli sites and the late Neolithic Liangchengzhen site (Figure 6).
Contextual variation is far greater at Liangchengzhen than at
either Zhangmatun or Xihe, but the mix of samples at the sites
includes a significant representation from pits and general cul-
tural layers. For the sake of hypothesis building, the samples are
roughly comparable and likely secondary or tertiary deposits; that
is, the remains were not recovered from contexts in which they
were stored, processed, eaten, or otherwise used. The Xihe per-
centages in Figure 6 are exaggerated because unidentifiable and
unknown seeds are not included in the site total, another factor
that compromises a nuanced comparison. Finally, taxa diversity is
impacted by sampling strategy, particularly sediment sample vol-
ume (Crawford et al., 2005). Flotation sampling was limited to a
small cross-section of sediments in each study. Sediment sample
sizes are mainly less than 10 L. Until larger samples are collected
and sampling strategies are designed to enable effective compari-
sons, we will not be able to adequately test hypotheses based on
such comparisons. Nevertheless, we feel that it is crucial to
explore such comparisons for future testing.
The crop percentages are similar at Xihe, Yuezhuang, and
Liangchengzhen, but crops are a minor component of the Zhang-
matun assemblage. This is because rice was not recovered from
Zhangmatun. Furthermore, foxtail millet comprises most of the
millet at Liangchengzhen while only a few broomcorn millet
grains have been recovered (Crawford et al., 2005). At both
Yuezhuang and Zhangmatun, broomcorn millet is the more com-
mon millet, although only eight millet grains were recovered from
Zhangmatun; six are broomcorn millet (Wu et al., 2014). Only
two millet grains (foxtail) are reported from Xihe (Jin et al.,
2014). Potential crops such as soybean, perilla, and chenopod are
components of all the assemblages.
Arboreal taxa represented at each site are notable for their dif-
ferences (only grape was recovered from all four sites). Arboreal
taxa are rare except at Zhangmatun where six taxa are reported,
grape being the most abundant (Wu et al., 2014). These differ-
ences may represent local preferences, specific habitat differ-
ences, sampling, or a combination of factors. The most common
arboreal plant seed at Liangchengzhen is a shrub/tree in the
legume family (Senna nomame) whose use is not clear (Crawford
et al., 2005). A large number of its seeds were found mainly in one
Figure 6. Comparison of Zhangmatun, Yuezhuang, Xihe, and
Liangchengzhen seed assemblages. The pie charts indicate sampled
context types. The bar chart indicates the percentage of significant
recovered plant taxa.
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10 The Holocene
sample. Arboreal plant fruit production is relatively high in open
habitats, so these taxa are still consistent with the hypothesis that
most of the plants were colonizing or possibly encouraged to
grow in anthropogenic habitats.
To what extent was the agricultural niche developing at
Yuezhuang and was it already similar to the Liangchengzhen
niche? First of all, the Zhangmatun, Xihe, Yuezhuang, and
Liangchengzhen assemblages are replete with small grass seeds
that appear to be mainly Paniceae, the tribe that also includes fox-
tail and broomcorn millet. They comprise a greater proportion of
the Liangchengzhen assemblage (particularly crabgrass, foxtail
grass type, and panic grass type), while the relative abundance
(percentage) of these grasses is similar at Zhangmatun, Xihe, and
Yuezhuang. Harlan describes swarms of weedy and wild grasses
in Africa growing among the millets (Harlan, 1989). Similarly,
swarms of grasses may have been developing along with culti-
vated millets in the Houli period and expanding their populations
in the late Neolithic as a result of increased disturbance. In other
words, as agriculture evolved so did weed communities. We do
not know at this time what types of grasses comprise the Paniceae
at Zhangmatun, but a similar range of types appears to be in the
Yuezhuang, Xihe, and Liangchengzhen assemblages, including
forms closely resembling Setaria sp. and Panicum sp. A few barn-
yard grass seeds have been recovered from each site too. The
Neolithic weed communities were likely developing by c. 9000–
8000 cal. BP.
Geoarchaeological research at Yuezhuang complements the
view that anthropogenic activities, including cultivation, were
crucial to the Houli Culture adaptation. The geophysical and geo-
chemical characteristics of the sediment provide further evidence
that cultural activities on the floodplain included clearing and
farming (Zhuang et al., 2013). Continuing sedimentation during
the occupation also likely facilitated plant cultivation over the
lifespan of the Yuezhuang occupation. Oxbow lakes and other
wetland environments were probably common because of the
evolution of the Yellow River drainage system (Zhuang et al.,
2013). Wetland animals were certainly important here (Song,
2003) as rice, too, appears to have been. Human agency was not
occurring in isolation. The extent to which post-Pleistocene
warming and the development of the Holocene landscape contrib-
uted to the development of subsistence economies in the Yellow
River valley needs further investigation. By 8000 cal. BP, the
Holocene Thermal Maximum (HTM) had started and may have
begun by 9000 cal. BP, preceded by a period of more intense mon-
soons (Renssen et al., 2012). Yuezhuang is situated at the northern
limit of the summer monsoonal zone today. The short-lived 8.2
cal. kyr BP event is also evident in China (Cheng et al., 2009).
Niche construction at the onset of the Neolithic must be set in this
environmental context.
Conclusion
The Yuezhuang data inform the general subsistence economy not
long after food production was established in Shandong Province.
Yuezhuang represents a relatively mature Houli Culture niche that
had a mixed economy of wetland exploitation, anthropogenic habi-
tat use, resource management, cultivation, hunting, gathering, and
fishing. The Houli presence on the landscape facilitated new eco-
logical opportunities and habitats for plant and animal populations,
some of which involved domestication that was well under way by
8000 cal. BP. Rice may have been introduced to the region after the
Zhangmatun occupation but before or during the Yuezhuang occu-
pation. Foxtail millet is a domesticated phenotype but relatively
rare at all Houli Culture sites. The conditions may not have been
conducive to it being an important crop. Instead, broomcorn millet
is common at Yuezhuang and six of the eight millet specimens
at Zhangmatun are broomcorn millet. No broomcorn millet was
identified at Xihe. Better sampling may help sort out the reasons for
the variable quantities of broomcorn millet. The preference for
broomcorn millet at Yuezhuang suggests that the crop was develop-
ing its relationship with people and was well along the path to
domestication. Finally, human activities related to food production
were beginning well before the Yuezhuang occupation. Evidence
includes the anthropogenic habitats evident as early as 9000 cal. BP
at Zhangmatun, the weed complexes that appeared at the same time
and became more established at Yuezhuang, and the presence of
crops and several plants that would eventually become crops.
Yuezhuang had a mixed economy that presages later Neolithic
economies in the region. The plant data from Yuezhuang combined
with the data from Zhangmatun and Xihe make the Houli Culture
arguably the best palaeoethnobotanical case study for the early
Neolithic in North China.
Acknowledgements
The analysis was conducted while Gary Crawford was the Liq-
ing Fellow in Social Sciences at Shandong University in April,
2004. Students from Shandong University collected and floated
the samples. Alison Dias composed the final version of Figure 1
and completed Figure 3. Liye Xie helped with translations of the
Chinese language literature.
Funding
The analysis was supported by the Liqing Foundation and the
follow-up research was facilitated by a grant from the Social Sci-
ences and Humanities Research Council of Canada (SSHRC
award 72022345). Support for the Yuezhuang project was also
obtained from the Youth Foundation of Humanities and Social
Science, Ministry of Education of China, 2011, the Independent
Innovation Foundation of Shandong University, and The Disci-
pline Innovative Engineering Plan.
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