Journal of Archaeological Science: Reports 51 (2023) 104152

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Journal of Archaeological Science: Reports

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Starch grains on human teeth as evidence for 4000 BCE potato consumption
at the Cruz Verde site, northern coast of Peru
Kazuho Shoji a, b, *, Víctor F. Vásquez S. c, Teresa E. Rosales T. d
a
Japan Society for Promotion of Science, Tokyo, Japan
b
Department of Humanities and Social Sciences, Yamagata University, Yamagata, Japan
c
Andean Archaeobiological and Paleoecological Research Center (ARQUEOBIOS), Trujillo, Peru
d
Archaeology program, Department of Social Science, University of Trujillo, Trujillo, Peru

A R T I C L E I N F O A B S T R A C T

Keywords: Cruz Verde, located on the north coast of Peru in the Chicama River Basin, is an artificial mound formed by the
Starch grains continuous disposal of food residues between 4200 and 3800 cal. BC. It is a vital archaeological site for un­
Dental calculus derstanding the early maritime community that was established in the coastal areas of Peru during the middle
Early plant use
Holocene period. Previous studies focusing on animal remains have shown that a wide range of marine resource
Maritime community
South America
uses were flexibly maintained in a changing environment. However, the use of plants by coastal fishermen, which
has been reported in recent years at other sites in the same region, was not evident at Cruz Verde. This paper
reports on an examination of starch grains preserved in the dental calculus of human teeth excavated from the
mound at Cruz Verde, which provides direct evidence of the early consumption of cultivated food plants, such as
maize (Zea mays), manioc (Manihot esculenta), squash (Cucurbita sp.), and potatoes (Solanum tuberosum). Not only
did the results indicate early use of maize indicated in previous studies, but they also confirmed that the use of
potatoes at the site dates back to at least 4000 cal. BC—the oldest example from the coastal region of Peru. The
micro-botanical data from Cruz Verde suggest that fishing and plant cultivation were combined in the devel­
opment process of early settlement. Furthermore, the data on this earliest potato use in the coastal area are
significant because they contribute to the empirical foundation of understanding the early relationship with
maritime and highland communities during this period.

1. Introduction
The establishment of a sedentary maritime community during the
middle Holocene period has been theorized as a key factor in recon­
structing the formation process of a complex society in the coast of
Central Andes (Moseley, 1975). Although many criticisms of this hy­
pothesis have been submitted over the stability of marine resources
(Osborn, 1977; Quilter, 1992; Quilter and Stocker, 1983) and the eval­
uation of phytophagy in subsistence economies (Haas and Creamer,
2006; Raymond, 1981; Wilson, 1981), the hypothesis has been revised
and the issues refined based on research in recent years (Beresford-Jones
et al., 2018; Sandweiss, 2009; Arriaza, 2008; Feldman, 2009), and its
influence is still significant today. One of the issues is the importance of
plant use by maritime communities (cf. Prieto and Sandweiss, 2020),
and recent archaeological researches are revealing that cultivated food
plants played an important role along with marine resources in early
costal sites (e.g. Benfer, 2008; Pearsall, 2008; Grobman et al., 2012;
Beresford-Jones et al., 2015, Beresford-Jones et al., 2021; Mauricio,
2015; Bonavia et al., 2017; Gorbahn, 2020; Tung et al., 2020; Dillehay
et al., 2022). Although flotation and fine screen sampling are the most
effective methods for obtaining macro-botanical remains (see Table 5),
there is expected to be variation in the persistence of these remains
depending on plant species, parts and burial environment. Therefore,
sampling of macro remain should be supplemented and verified by
micro-botanical sampling following a solid methodology to understand
the relationship between marine resource and plant resource use in the
development process of early sedentary settlement.
In the Chicama River basin, several preceramic mounds with abun­
dant natural remains are distributed along the coastline (Bird et al.,
1985; Dillehay et al., 2012; Dillehay, 2017; Hirota, 2008; Maggard and
Dillehay, 2017). One of the sites, Cruz Verde, is located on a small
coastal terrace approximately 200 m from the coastline, 3 km north of

* Corresponding author.
E-mail address: lpo.spd1418@gmail.com (K. Shoji).

https://doi.org/10.1016/j.jasrep.2023.104152
Received 17 April 2023; Received in revised form 25 July 2023; Accepted 25 July 2023
2352-409X/© 2023 Elsevier Ltd. All rights reserved.
K. Shoji et al.
Huaca Prieta and Paredones, and 12 km south of Huaca Pulpar (Fig. 1).
The excavation results revealed that the formation process of the Cruz
Verde mound can be divided into CV-Ia (4200–4000 cal. BC) and CV-Ib
(4000–3800 cal. BC) phases, and the repetitive overlapping of floor
surfaces and sedimentary layers encompassing faunal remains were
continuously present throughout the two phases (La Rosa and Shoji,
2017, 2018; Shoji, 2021). Especially from the CV-Ib phase, nine human
burials were recovered from the mound, and distinct floors made of clay
were found. The excavated fauna also changed at the same time (Shoji,
2018, 2022). This indicates that the processes of mound formation and
resource use changed significantly during the two phases. All the fauna
excavated were from the marine environment, suggesting a strong
dependence on marine resources; however, there were also many arti­
facts such as grinding stones and stone plates that suggest the use of
plants. Therefore, a micro-botanical analysis was conducted to confirm
this.
2. Materials
We examined the teeth of five of the nine excavated burials that were
in a good state of preservation (Fig. 3). Human remains were buried in a
variety of ways and at different times of mound formation. Some were
placed in an open pit cutting into the floor or sedimentary layer, some
were embedded in the floor and covered by a sedimentary layer, and
some were embedded during the sedimentation process; no fixed pat­
terns in burial posture were observed (Shoji, 2021). There were very few
funeral offerings, except for one simple polychaete tube-base figurine
(cf. Dillehay and Bonavia, 2017), and all the burials were simply
wrapped in plant mats. And there is no pattern of burial structure,
cluster, or burial position, which is often used as archaeological evi­
dence to discuss social class differences among burials, in addition, there
are no differences in pathological features. However, it is possible that
certain groups within the maritime community were interred at the
Fig. 1. Location map of Cruz Verde and other sites discussed in this study.
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Journal of Archaeological Science: Reports 51 (2023) 104152
mound, as all of the burials were detected as adults and eight of the nine
human remains were male (Palma, 2019). Each burial was simply
wrapped in mat-like material made of woven plants, and accelerator
mass spectrometry (AMS) dating was performed on the plant remains
attached to the human remains (Shoji et al., 2018) (Fig. 3). Their ages
fall within the range of 4000–3800 cal. BC and are consistent with
stratigraphic data (Table 1). Because the human remains from Cruz
Verde, a community that is heavily marine-derived, are not suitable for
accurate dating due to the significant influence of marine reservoirs,
these plant remains, which are unquestionably directly related to burial,
were used as samples for dating.
Thick dental calculus deposits were observed on almost all teeth in
the five well-preserved burials (Fig. 2a), suggesting that the burials were
generally in an oral environment prone to dental calculus deposits (cf.
Tung et al., 2020). Thick deposits were especially common on molars,
and we obtained one to three tooth samples from each individual.
Although no previous systematic studies on the rate of dental calculus
formation are available, it is assumed that each tooth reflects at least
several years of diet, as calculus will accumulate over a lifetime, if not
removed (Piperno and Dillehay, 2008).
Similarly, for lithic tools on which starch grains were examined,
organic material was collected from the 2–3 mm diameter depressions
and fractures that remained on the surface with use-wear of grinding
stone and stone plate. Of the seven lithic artifacts, four were selected
from the sedimentary layers, and the remaining three were reused to
support the burial posture placing under the body (Fig. 4). One undec­
orated gourd container, which was also placed like a pillow with the
opening facing the ground under the head of burial TM10, was selected,
and the organic material adhering to its interior walls was collected.
K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Table 1
Radiocarbon dates on plant mat wrapping human burials and wood charcoal from stratigraphic layer in excavated unit of Cruz Verde.
14
Lab No. Registry No. Provenience Material C Age δ13C (‰) Calibrated age Calibrated age
(BP) range range
(cal BC) 1σ (cal BC) 2σ

Radiocarbon dates for human

burials
TKA-21441 16CV2-A- Unit SV1, Plant 5029 ± 24 − 27.5 ± 3790–3710 3935–3875
P048 Layer 7a, Plant mat Mat 0.4 (68.2%) (13.4%)
wrapping TM5 3810–3690
(80.1%)
3685–3731
(2.0%)
TKA-18006 16CV2-A- Unit 1701, Grid O1N5, Plant 5036 ± 28 − 24.4 ± 3890–3885 3940–3855
C045 Layer 12a, Plant mat wrapping Mat 0.6 (2.1%) (21.4%)
TM7 3800–3705 3815–3690
(66.1%) (72.3%)
3685–3660
(1.7%)
PLD-36344 17CV-A- Unit 1701, Grid O2N4, Surface of Plant 5174 ± 23 − 10.0 ± 3980–3940 4035–4025
C010 Layer 12a, Mat 0.2 (44.0%) (0.7%)
Plant mat wrapping TM8 3860–3815 3995–3910
(24.2%) (56.3%)
3880–3800
(38.4%)
PLD-36345 17CV-A- Unit 1701, Grid O1N4, Plant 5088 ± 23 − 26.3 ± 3945–3855 3955–3765
C020 Layer 12a, Plant mat wrapping Mat 0.2 (52.4%) (95.0%)
TM9 3820–3785 3725–3715
(15.8%) (0.4%)
TKA-21442 17CV-A- Unit 1701, Grid O2N4, Plant 5411 ± 26 − 18.9 ± 4325–4285 4330–4220
P068 Layer 12e, Plant mat wrapping Mat 0.6 (18.4%) (53.1%)
TM10 4270–4225 4215–4150
(27.1%) (22.4%)
4205–4165 4135–4055
(16.7%) (19.9%)
4130–4120(1.7%)
4095–4080
(4.3%)
Radiocarbon dates for
stratgraphic layeres
PLD-36349 17CV-A- Unit 1701, Grid E2S1, Wood 5111 ± 24 –22.5 ± 3945–3910 3960–3790
C052 Layer 8a Charcoal 0.2 (20.1%) (95.4%)
3880–3800
(48.1%)
TKA-18005 16CV2-A- Unit 1701, Grid O1S1, Carbon Wood 5296 ± 22 − 30.5 ± 4225–4205 4230–4195
C037 Layer 12b Charcoal 0.7 (5.9%) (10.6%)
4155–4130 4175–4090
(12.0%) (26.2%)
4065–3985 4085–3975
(50.3%) (58.6%)
PLD-36341 16CV2-A- Unit 1701, Grid O1S1, Carbon Wood 5279 ± 22 − 28.0 ± 4055–3980 4230–4200
C040 Layer 12d Charcoal 0.1 (68.2%) (5.4%)
4170–4125
(10.5%)
4115–4095
(2.0%)
4080–3965
(77.4%)
PLD-36340 16CV2-A- Unit 1701, Grid O1S1, Surface of Wood 5292 ± 21 − 28.5 ± 4225–4210 4230–4200
C028 layer 15a Charcoal 0.2 (4.5%) (9.2%)
4155–4130 4170–4090
(9.7%) (22.2%)
4060–3985 4085–3975
(54.0%) (63.9%)
TKA-18918 16CV2-A- Unit 1701, Grid O1S1, Surface of Wood 5380 ± 28 –22.6 ± 4245–4220 4320–4295
C041 natural Charcoal 0.3 (9.6%) (3.4%)
ground layer 4210–4155 4265–4045
(27.1%) (92.0%)
4135–4065
(31.5%)

*All dates in Table 1 were calibrated using SHcal13 (Hogg et al., 2013).

3. Methods (Piperno and Dillehay, 2008), employing nondestructive methods for

teeth and starch grains to remove dental calculus and examine the starch
3.1. Starch grain sampling of teeth in it. From five well-preserved human remains that had been processed
through physical anthropological analysis, one to three incisors and
We followed the sampling methods established in previous studies molars with dental calculus were selected.

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K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Fig. 2. (a) A human molar with thick dental calculus deposit recovered from a human burial at Cruz Verde. (b) A faceted starch grain from Cucurbita sp., extracted
from CV-AL-13 (grinding stone). Left, non-polarized view. Right, polarized view. (c) A hemispheric starch grain from Cucurbita sp., extracted from CV-AL-07
(grinding stone). Left, non-polarized view. Right, polarized view. (d) A polyhedric starch grain from Zea mays with a fissure on hilum, extracted from CV-AL-02
(human burial, TM7). Left, non-polarized view. Right, polarized view. (e) A polyhedric starch grain from Zea mays, extracted from CV-AL-03 (human burial,
TM8). Left, non-polarized view. Right, polarized view. (f) A truncated bell shape starch grain from Zea mays, characteristic of the “morocho” race, extracted from CV-
AL-09 (stone plate). Non-polarized view. (g) An elliptical starch grain from Solanum tuberosum, extracted from CV-AL-03 (human burial, TM8). Left, non-polarized
view. Right, polarized view. (h) A trapezoidal starch grain from Solanum tuberosum, extracted from CV-AL-16 (gourd container). Non-polarized view. (i) A hemi­
spheric starch grain from Manihot esculenta, extracted from CV-AL-13 (grinding stone). Left, non-polarized view. Right, polarized view.

First, the tooth was lightly washed with a soft toothbrush and
distilled water to remove soil and other debris. A disposable razor was
used to remove a thin surface layer of the calculus, after which a new
razor was used for further inner calculus sampling. The razor was
replaced with a new one for each sample. The collected calculus was
placed in a test tube with distilled water, and kept for five days with slow
agitation, until homogenization. One drop of this was then transferred to
a slide, and a 5% saline solution and glycerin were added to retard
drying and facilitate rotation of the grains as they were encountered.
Glycerin did not alter or damage the morphology of the starch grains.
Coverslips were applied and microscopic observations were made.
3.2. Starch grain sampling of lithic tool and gourd
Regarding the lithic tools, the excavated grinding stones and stone
plates were selected and cleaned using a soft toothbrush and compressed
air duster, and organic material was collected from the depths of the 2–3
mm depressions and cracks that remained on the surface of use (Fig. 4).
When samples were collected, a low-magnification microscope was used
to confirm that it was a primary sedimentary layer of organic material,
and a new needle was used after the thin surface layer was removed. The
samples were stored for five days under agitation in distilled water,
similar to the calculus samples.
Samples of the organic material adhering to the interior of the gourd
container were collected using a new disposable razor after the thin
surface layer had been removed. One drop from each sample was
transferred to a glass slide, and microscopic observations were made
using the same procedure as for the calculus described earlier.
3.3. Starch grain identification
Identification was performed using modern reference collections,
mainly of edible native plants, archived at ARQUEOBIOS, as well as
other published sources (Pagán, 2015; Piperno and Dillehay, 2008;
Torrence and Barton, 2006; Loy, 1990; Piperno, 2006). The starch
classification system we used emphasizes basic morphological data,
such as shape, position of the hilum, facets, and fissure patterns, which
are known to be useful for identification (Fig. 2) (Piperno, 2006).
A polarizing microscope was used to observe and record the external
shape of the starch grains, the shape and size of the polarizing crosses at
400x magnification. Polarized light is typically used to detect starch
bodies. Under polarized light, starch grains show an interference figure
in the form of a “cross,” which is the result of the double refraction of
light through the cylindrical or spherical lamellar structures of the
starch grains. The point of intersection between the two parts of the
cross generally corresponds to the hilum position. This is an opening or
cross-section of the tube or passage into the interior of the grain, through
which the starchy matter forms the internal lamina that passes. Thus, a
microscopic examination of the starch grains with a polarizing micro­
scope is unequivocal evidence that its double refractive power can be
obtained and its presence confirmed. Measurements were taken of the
identified and unidentified and/or damaged starch grains using a
measuring device or reticule calibrated to microns, which was attached
to the eyepiece of the microscope. The maximum length and width of
each identified and unidentified starch grain were measured.
4. Results

Significant amounts of starch grains were found in most of the

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K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Fig. 3. Human burials and characteristic microbotanical remains from dental calculus at Cruz Verde. Dental calculus of good preserved five human remains (TM3
and TM7-10) are analyzed. See Table 3 for more detail information of identified micro remains.

human teeth collected from the five burials. Phytoliths were also found
in small quantities. Table 2 shows the types and number of starch grains
collected from these samples. Zea mays was detected in all five in­
dividuals, followed by Cucurbita sp. and Solanum tuberosum in three in­
dividuals (Table 2, 3). These three types of starch grains were also
detected on the stone tools and the gourd container (Table 2, 4), along
with a small amount of Manihot esculenta. The fact that starch grains
biased toward the above four taxa were equally detected on the pro­
cessing tools and on human teeth indicates that these plant species were
processed and consumed as major food plants.
The presence of Zea mays, in which an especially large number of
starch grains were detected, has been confirmed in the dental calculus
from TM10, the oldest burial at the end of the CV-Ia phase, as well as in a
grinding stone (CV-AL-10) from the sedimentary layer of the CV-Ia

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K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Fig. 4. Analyzed stone tools and characteristic micro-botanical remains recovered at Cruz Verde. White triangle on the stone tools indicate points where organic
remains were collected. See Table 4 for more detail information of identified micro remains and archaeological context.

phase and two grinding stones recovered in association with TM10 (CV-
AL-07, 08) (Fig. 4 and Table 4). The detection of Zea mays starch grains
on all CV-Ia and CV-Ib lithic tools indicated that the use of maize, in
particular, continued to be prominent from the beginning of the occu­
pation. The starch grains found on the stone plate in the current study
(CV-AL-09: CV-Ib phase) show a truncated bell shape characteristic of
“morocho,” an old race of maize (Fig. 2f). The starches of this maize race
are generally simple and quite irregular in which several variants of
truncated forms predominate. Of these, the most common shape is the
truncated with a widened proximal section, followed by the elongated
and narrow truncated form (Pagán, 2015). This proto-race has also been
recovered as macro-remains at other preceramic coastal sites, such as
Los Gavilanes and Paredones (Grobman et al., 2012; Bonavia, 1982).
Starch grains with hemispheric, ovoid, and faceted shapes

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Table 2
Types and number of starch grains recovered from Cruz Verde.
Sample no., calibrated age (cal BC/ Material, Z. mays Cucullbita S. tuberosum M. esculenta No TOTAL
2σ) asociation sp. Id.

Phase n n n n n n
Dental calculus
CV-AL-04, 3955–3715, PM DC, TM9 CV-Ib 4 2 1* 7
CV-AL-01, 3960–3790, WC DC, TM3 CV-Ib 7 3 4 14
CV-AL-02, 3960–3660, PM DC, TM7 CV-Ib 5 1 6
CV-AL-03, 4035–3800, PM DC, TM8 CV-Ib 8 1 3 12
CV-AL-05, 4330–4055, PM DC, TM10 CV-Ia 5 1* 6
Stone tools and Gourd
container
CV-AL-09, 3960–3790, WC SP, L8a CV-Ib 6 1 7
CV-AL-11, 3955–3715, PM GS, L12a CV-Ib 5 5
CV-AL-12, 3955–3715, PM GS, L12a CV-Ib 7 2* 1 1* 11
CV-AL-13, 3955–3715, PM GS, TM9 CV-Ib 3 2 1 2 8
CV-AL-10, 4230–3965, WC GS, L12d CV-Ia 4 1 5
CV-AL-16, 4330–4055, PM GC, TM10 CV-Ia 5 4 9
CV-AL-07, 4330–4055, PM GS, TM10 CV-Ia 7 1 2* 10
CV-AL-08, 4330–4055, PM GS, TM10 CV-Ia 7 7

Note: Calibrated ages for all samples are derived from materials (PM = plant mat wrapping human remains, WC = wood charcoal associated with carbon layer or
human burial) directly associated with teeth or artifacts in an archaeological context, sealed in individual intact floors (see Table 1). The material of each sample is
indicated as DC = dental calculus, SP = stone plate, GS = grinding stone, GC = gourd container. TM indicates the registration number of human burials and L indicates
the number of layers associated with samples (see also Fig. 3 and 4). *Taxa Identified as characteristic phytoliths.

characteristic of Cucurbita sp. were found on CV-Ia stone tools (Table 2,
4). In addition, phytoliths, which are found in squash peels, have been
detected in dental calculus and grinding stones, suggesting that the
whole fruit was consumed with its peel. Seeds of C. moschata were
recovered from the sedimentary layers of the Cruz Verde site (Table 5);
however, the feature of some truncated, bell-shaped starch grains with
measurements between 15.6 and 20 µm indicates that they may be
C. ficifolia. Nevertheless, the results confirm previous studies on moun­
tainous sites, suggesting that these cultivated plants were also consumed
as a food resource in the early period (Piperno and Dillehay, 2008;
Rossen, 2011), even in coastal areas. Although the number of starch
grains and phytoliths of Cucurbita sp. was not as high as that of Zea mays,
Cucurbita pulp should be considered to have been consumed continu­
ously since the CV-Ia phase, even considering its low starch production
(Piperno and Dillehay, 2008).
Starch grains of Solanum tuberosum collected from dental calculus
provide important evidence for reconsidering the history of potato use in
coastal areas. Potatoes are thought to have been cultivated in the An­
dean altiplano, and the only evidence from the coastal region was re­
ported as the macro-botanical remains found at a few sites, such as
Huaynuna in the Casma Valley, dated approximately 2000 BCE (Ugent
et al., 1982; Ugent and Peterson, 1988). The starch grains of Solanum
tuberosum detected from three individuals, the first micro-botanical re­
mains reported on the coast, showed elliptical and trapezoidal shapes,
with a hilum in an eccentric position that is characteristic of the taxa
(Fig. 2g–e). Its size was somewhat smaller than that of those grown in
the Andean altiplano ecosystem, which may be the result of adaptation
to the coastal ecosystem. The fact that stone tools and burials of TM3,
TM8, and TM9, in which Solanum tuberosum starch grains were detected,
all belong to CV-Ib indicates that potatoes were completely consumed in
this phase (Table 1, 2). Although Solanum tuberosum starch grains were
not detected on stone tools in the CV-Ia phase, a significant number of its
starch grains were also detected on the inner surface of a gourd
container that was associated with the human remains of TM10– the
oldest burial belonging to the end of CV-Ia (Table 1, 4 and Fig. 3). Based
upon these data, we can say that the use of potatoes at Cruz Verde
occurred at the end of the CV-Ia period, around 4000 cal. BC, at the
latest. However, the number of Solanum tuberosum starch grains found
on stone tools of CV-Ib was not very high, which may indicate that these
tools were not used much for potatoes even in this phase.
The Manihot esculenta starch grains collected from two grinding
stones in the CV-Ib phase also represent a new micro-botanical remain
identified at a preceramic coastal site. Although only one starch grain
was identified from each lithic and none was detected in the dental
calculus, the hemispheric shape and size are characteristic of Manihot
esculenta (Fig. 2i). As there are some slightly toxic varieties of manioc, it
is possible that tubers were grounded to remove the toxin. The use of
manioc in coastal areas, first found at the Cruz Verde site, is thought to
date back to 4000–3800 cal. BC.
5. Discussion and conclusions
5.1. Plant use by maritime community
Data on starch grains collected from the dental calculus, stone tools,
and gourd container indicate that the preceramic coastal fishermen used
food plants that could not be detected in macro-botanical remains ob­
tained from soil flotation samples and other sources (Table 5). Similar
use of maize and other plants by costal people at the nearby Huaca Prieta
and Paredones sites has been reported in recent years (Grobman et al.,
2012; Bonavia et al., 2017; Tung et al., 2020), especially in the case of
Paredones, which is slightly older than the data from Cruz Verde, indi­
cating that the use of maize in this region started before Cruz Verde
(Grobman et al., 2012; Vallebueno-Estrada, 2023). The prominent Zea
mays starch grains and macro-botanical data from Cruz Verde may
indicate part of a long-term process of introduction and establishment of
such plant use. This clearly supports the argument of Grobman et al.,
that maize utilization in the coastal areas of the Chicama River Basin
dates back to 4825–4554 cal. BC (Grobman et al., 2012). The fact that
these data were obtained from Cruz Verde, where marine resources were
abundantly exploited (Shoji, 2018, 2022), re-affirms that a combined
fishing and main crops farming economy was established on the north
coast during this period (cf. Dillehay et al., 2022).
While there are similarities in the use of maize by these coastal
fishermen with Huaca Prieta and Paredones, there are also some dif­
ferences in the detection of micro-botanical remains. At Huaca Prieta, in
addition to the analysis of starch grains in the soil, the dental calculus of
nine human burials excavated in 2009 and nine collected by Bird were
analyzed. Data from the 2009 excavation showed that only two of the
nine samples had starch grains in their dental calculus; of these, one
sample could not be identified, and the other had only one starch grain
from Phaseolus sp., which could be a lima bean (Piperno et al., 2017). As
for the human remains collected during excavations by Bird, starch
grains of Phaseolus sp. and Cucurbita moschata were found in three of the

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Table 3
Identified taxa, size and radiocarbon dates of all micro botanical remains extracted from dental calculus of five human remains at Cruz Verde.
Sample no. Burial no. Calibrated age Dentition no. Taxon Length £ Width (μm) Type of remains Shape, Description
(BC, 2σ), Source

CV-AL-01 TM3 3960–3790, WC #26 Zea mays 18.2 × 15.6 Starch grain polyhedric
(lateral incisor) 18.2 × 18.2 Starch grain polyhedric
#31 20.8 × 20.8 Starch grain polyhedric, fissure on hilm
(second molar) 15.6 × 13 Starch grain polyhedric
15.6 × 15.6 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
15.6 × 13 Starch grain polyhedric, fissure on hilm
Cucurbita sp. 10.4 × 10.4 Starch grain faceted
11.7 × 10.4 Starch grain ovoid
11.7 × 10.4 Starch grain faceted
S. tuberosum 18.2 × 13 Starch grain elliptical
15.6 × 15.6 Starch grain trapezoidal
16.9 × 13 Starch grain trapezoidal
13 × 10.4 Starch grain ovoid
CV-AL-02 TM7 3960–3660, PM #25 Zea mays 18.2 × 18.2 Starch grain polyhedric
(third molar) 20.8 × 18.2 Starch grain polyhedric, fissure on hilm
#26 18.2 × 18.2 Starch grain polyhedric
(central incisor) 20.8 × 20.8 Starch grain polyhedric
#32 18.2 × 13 Starch grain polyhedric
(lateral incisor)
Cucurbita sp. 11.7 × 11.7 Starch grain faceted
CV-AL-03 TM8 4035–3800, PM #17 S. tuberosum 10.4 × 10.4 Starch grain spherical
(lateral incisor) 13 × 13 Starch grain spherical
#23 26 × 20.8 Starch grain elliptical
(third molar)
Cucurbita sp. 13 × 13 Starch grain faceted

Zea mays 20.8 × 18.2 Starch grain polyhedric

18.2 × 13 Starch grain polyhedric
14.3 × 14.3 Starch grain polyhedric
18.2 × 18.2 Starch grain polyhedric
18.2 × 16.9 Starch grain polyhedric
14.3 × 14.3 Starch grain polyhedric
19.5 × 13 Starch grain polyhedric
18.2 × 16.9 Starch grain polyhedric
CV-AL-04 TM9 3955–3715, PM #24 S. tuberosum 28.6 × 23.4 Starch grain ovoid
(central incisor) 18.2 × 11.7 Starch grain elliptical

Zea mays 18.2 × 18.2 Starch grain polyhedric

14.3 × 14.3 Starch grain polyhedric
15.6 × 13 Starch grain polyhedric
15.6 × 13 Starch grain hemispheric
No Identified Phytolith
CV-AL-05 TM10 4330–4055, PM #19 Zea mays 20.8 × 18.2 Starch grain polyhedric
(first molar) 14.3 × 13 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
18.2 × 14.3 Starch grain polyhedric
Cucurbita sp. 31.2 × 23.4 Phytolith spherical scalloped

Note: Source of calibrated age are indicated as PM=plant mat or WC=wood charcoal (see Table 1). Dentition no. is indicated using universal numbering system for
teeth.

eight samples from which dental calculus was successfully collected. In
contrast, in the current Cruz Verde study, maize was identified in all
dental calculus, and no Phaseolus sp. was recovered at all, not even
macro-botanical remains (Table 5). The possibility cannot be ruled out
that the above results may be due to conditions of preservation or other
factors, as the number of starch grains detected in the dental calculus
from the Huaca Prieta is low, and also, there is a lack of dental calculus
data at Paredones, which is known to have consumed more maize than
Huaca Prieta (Tung et al., 2020). But at least, the absence of Phaseolus sp.
in the data from Cruz Verde, where many other starch grains were
detected, suggests that there may have been different plant use trends at
Cruz Verde and Huaca Prieta.
It is interesting to note that similar differences can be examined in
other data as well. The difference is that no maize and potato was
detected on a gourd container (4500–4400 cal. BC) from Paredones
(Phaseolus sp. and Inga feuillei were detected) in contrast to the same type
of material from Cruz Verde (Table 4). Furthermore, abundant starch
grains of maize were detected on grinding stones and plate from Cruz
Verde, whereas only Cucurbita sp. was detected on a grinding stone from
Huaca Prieta (Piperno et al., 2017). As mentioned above, many macro-
botanical remains of maize have been excavated from Huaca Prieta and
Paresones, and maize starch grains have been detected in the soil and on
the blades of the flaked stone tools. It also seems that the use of maize on
the north coast had already been introduced before the occupation of
Cruz Verde (cf. Dillehay et al., 2022; Grobman et al., 2012; Vallebueno-
Estrada, 2023). Thus, there is no doubt that maize was used at all three
sites, but the differences in the data obtained from the stone tools and
gourd container also suggest that there may have been differences in the
way it was used. However, these data are still a small sample size, and
the different archaeological contexts of the artifacts should also be taken
into consideration. Future data collection and verification is required to
understand the differences among the three sites. It should be added that
only one macro-botanical remain of chile pepper, which was excavated
in large quantities at Huaca Prieta and Paredones (Chiou et al., 2014),
was recovered from Cruz Verde (Table 5).

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K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Table 4
Identified taxa, size and radiocarbon dates of all micro botanical remains extracted from stone tools and gourd container at Cruz Verde.
Sample no. Regitration no. Calibrated age Provenience Type of artifact Taxon Length x Type of remains Shape, Description
of materials (BC, 2σ), Source Width (μm)

CV-AL-07 17CV-A-L095 4330–4055, PM Unit 1701, grinding stone Zea mays 13 × 13 Starch grain polyhedric
Grid O2N4, 18.2 × 15.6 Starch grain polyhedric
asociated with 15.6 × 15.6 Starch grain polyhedric
TM10 16.9 × 15.6 Starch grain polyhedric
13 × 13 Starch grain polyhedric
13 × 10.4 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
Cucurbita sp. 13 × 13 Starch grain hemispheric
Poaceae 78 × 15.6 Phytolith
No Identified 46.8 × 39 Phytolith
CV-AL-08 17CV-A-L118 4330–4055, PM Unit 1701, grinding stone Zea mays 18.2 × 18.2 Starch grain polyhedric
Grid O2N4, 18.2 × 15.6 Starch grain polyhedric
asociated with 18.2 × 16.9 Starch grain polyhedric
TM10 20.8 × 18.2 Starch grain polyhedric
15.6 × 15.6 Starch grain polyhedric
16.9 × 14.3 Starch grain polyhedric
26 × 18.2 Starch grain polyhedric
CV-AL-09 16CV2-A-L145 3960–3790, WC Unit 1701, stone plate Zea mays 15.6 × 15.6 Starch grain polyhedric, perforated
Grid O1N5, hilm
Layer 8a, hoyo3 18.2 × 18.2 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
15.6 × 15.6 Starch grain polyhedric
20.8 × 15.6 Starch grain polyhedric
15.6 × 13 Starch grain truncated, “morocho”?
cf.S. tuberosum 15.6 × 13 Starch grain trapezoidal
CV-AL-10 17CV-A-L043 4230–3965, WC Unit 1701, grinding stone Zea mays 15.6 × 15.6 Starch grain polyhedric
Grid O2N4, 20.8 × 20.8 Starch grain polyhedric
Layer 12d 19.5 × 19.5 Starch grain polyhedric
18.2 × 18.2 Phytolith phytolith of maize cob
No Identified 15.6 × 15.6 Starch grain ovoid
CV-AL-11 17CV-A-L058 3955–3715, PM Unit 1701, grinding stone Zea mays 18.2 × 15.6 Starch grain polyhedric
Grid O1S5, 14.3 × 13 Starch grain polyhedric
Layer 12a 13 × 13 Starch grain polyhedric
15.6 × 13 Starch grain polyhedric
15.6 × 15.6 Starch grain polyhedric
CV-AL-12 17CV-A-L059 3955–3715, PM Unit 1701, grinding stone Zea mays 23.4 × 20.8 Starch grain polyhedric
Grid O1S5, 18.2 × 18.2 Starch grain polyhedric
Layer 12a 20.8 × 20.8 Starch grain polyhedric
18.2 × 15.6 Starch grain polyhedric
16.9 × 16.9 Starch grain polyhedric
16.9 × 15.6 Starch grain polyhedric
18.2 × 16.9 Starch grain polyhedric
M. esculenta 18.4 × 18.4 Starch grain hemispheric
Cucurbita sp. 28.6 × 26 Phytolith spherical scalloped
44.2 × 36.4 Phytolith spherical scalloped
No Identified 23.4 × 10.4 Phytolith
CV-AL-13 17CV-A-L101 3955–3715, PM Unit 1701, grinding stone Zea mays 20.8 × 18.2 Starch grain polyhedric
Grid O1N4, 14.3 × 13 Starch grain bell shaped
asociated with 13 × 10.4 Phytolith phytolith of maize leaf
TM9
Cucurbita sp. 13 × 10.4 Starch grain polyhedric
13 × 10.4 Starch grain polyhedric
M. esculenta 23.4 × 23.4 Starch grain hemispheric
26 × 23.4 Starch grain hemispheric
S. tuberosum 13 × 11.7 Starch grain elliptical
CV-AL16 17CV-A-P68 4330–4055, PM Unit 1701, gourd container S. tuberosum 24.7 × 20.8 Starch grain elliptical
Grid O2N4, 13 × 13 Starch grain trapezoidal
asociated with 18.2 × 15.6 Starch grain elliptical
TM10 14.3 × 10.4 Starch grain elliptical
Zea mays 13 × 10.4 Starch grain polyhedric
13 × 13 Starch grain polyhedric
13 × 13 Starch grain polyhedric
20.8 × 18.2 Starch grain polyhedric
18.2 × 18.2 Starch grain polyhedric

Note: Source of calibrated age are indicated as PM=plant mat or WC=wood charcoal (see Table 1).

5.2. Significance of potato consumption
The data from Cruz Verde are also important because they reveal for
the first time the presence of Solanum tuberosum and Manihot esculenta
around 4000 cal. BC in the region. These two species, one tuber and
other root tuber, have not been identified at Huaca Prieta or Paredones,
and their provenance as macro-botanical remains has been limited to
2000 BCE (Ugent et al., 1982; Ugent and Peterson, 1988). This means
that the data for the two species from Cruz Verde is the earliest example
from a Peruvian coastal site. It is clear that the preceramic coastal
fishermen on the north coast of Peru used not only Zea mays and
Cucurbita sp., but also various plant crops, including Solanum tuberosum

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K. Shoji et al. Journal of Archaeological Science: Reports 51 (2023) 104152

Table 5 Data availability

Identified taxa and number of macro-botanical remains of flotation sample
excavated at Cruz Verde. Data will be made available on request.
Family Genus and CV-Ia phase CV-Ib phase Total
Species (4200–4000 cal (4200–4000 cal Acknowledgments
BC) BC)
n n n
This research was supported by the Keunokai Young Researcher
Poaceae Zea mays 2 0 2 Support Program, JSPS Grant-in-Aid for JSPS Fellows (Grant number
Malvaceae Gossypium 1 1 2
16J06775, 2016-2017), Takanashi Foundation for Arts and Archaeology
barbadense
Solanaceae Capsicum sp. 1 0 1 (2015 and 2017), Graduate University for Advanced Study (SOKENDAI)
Myrtaceae Psidium guajava 23 0 23 Student Dispatch Program (2015-2017), and Graduate University for
Psidium sp. 15 0 15 Advanced Study (SOKENDAI) Internship Program (2017). The Paleo
Cucurbitaceae Cucurbita 0 1 1 Labo Young Researcher Support Foundation funded the AMS analysis.
moschata
Lagenaria 1 0 1
The Laboratory of Radiocarbon dating in the museum of the University
siceraria of Tokyo and Takayuki Omori supported the dating analysis. We thank
Fabaceae Prosopis sp. 0 3 3 the Instituto Nacional de Cultura of Peru, especially Jesús Briceño, Flor
Acacia sp. 18 12 30 Diaz, and César Gálvez (INC, Trujillo). We are grateful to Martha Palma,
Cyperaceae Cyperus sp. 3 0 3
who analyzed human remains, Segundo Vásquez (UNT), José Samuel
Scirpus sp. 1 0 1
No identified 14 4 18 Querevalú, Pedro Caceres, and Vanessa La Rosa.
Total 79 21 100
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and Manihot esculenta, as well as marine resources. It is possible that the
resource use of the early sedentary maritime communities that emerged
from the middle Holocene period was established as a combination of
fishing and cultivation using coastal wetlands, as described by Dillehay
et al. (Dillehay, 2017).
Potatoes are highland tubers that are thought to be domesticated
multicentrically in the Andean altiplano, and the excavated remains at
the highland sites are older than those of Cruz Verde (Engel, 1970). It
seems reasonable to assume that domesticated potatoes were brought to
Cruz Verde from the highlands. However, there is no archaeological
evidence at Cruz Verde indicating interregional interaction with the
Andean highlands, except for the presence of potatoes. It is therefore still
not clear how the potato use was initiated in the coastal area. Further,
we should also point out that a wild costal species of potato, different
from the highland species, exists naturally in the Lomas vegetation of
Cerro Campana, approximately 30 km from Cruz Verde (Ochoa, 1999;
Dillon, 2005). And the wild species, Solanum mochiquense, had been
collected and consumed by local people near Cerro Campana (Ochoa,
1999). Therefore, how coastal preceramic potato use was initiated must
be clarified by genetic analysis of potato samples from Cruz Verde,
compared to wild potato species from coastal and highland regions.
Sedentary groups that exploited marine resources during this period
have been given an important place as the basis for the construction of
monumental architecture and complex ideological, technological, and
political developments that began after 3500–3000 BCE (Moseley, 1975;
Feldman, 1985; Shady and Leyva, 2003; Chu, 2011; Mauricio, 2015).
The data from Cruz Verde, as other similar data show (e.g. Bonavia et al.,
2017; Piperno et al., 2017; Tung et al., 2020), indicate that the actual
use of such resources was not simply limited to marine resources, and a
reexamination of the economic activities and social organization of
these communities is warranted.
Author Contributions.
Kazuho Shoji conceived the idea of the study. Víctor F. Vásquez and
Teresa E. Rosales conducted micro-botanical remain sampling and plant
species identification. Kazuho Shoji contributed to the interpretation of
the results and drafted the original manuscript. All authors reviewed the
manuscript draft and revised it critically on intellectual content. All
authors approved the final version of the manuscript to be published.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
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