ORIGINAL RESEARCH
published: 20 September 2019
doi: 10.3389/fpls.2019.01161
Hidden Rice Diversity in the Guianas
Tinde Van Andel 1,2*, Margaretha A. Veltman 3, Alice Bertin 1, Harro Maat 4, Thomas Polime 5,
Derk Hille Ris Lambers 6, Jerry Tjoe Awie 7, Hugo De Boer 1,3 and Vincent Manzanilla 3
1 Department Biodiversity Dynamics, Naturalis Biodiversity Center, Leiden, Netherlands, 2 Biosystematics group, Wageningen
University, Wageningen, Netherlands, 3 Natural History Museum, University of Oslo, Oslo, Norway, 4 Knowledge,
Technology and Innovation group, Wageningen University, Wageningen, Netherlands, 5 Wooko Makandie Foundation,
Culemborg, Netherlands, 6 Dassenboslaan 15, Wageningen, Netherlands, 7 Department Research Management & Plant
Breeding, Anne van Dijk Rice Research Centre (SNRI/ADRON), Nieuw Nickerie, Suriname
Edited by:
Nina Rønsted,
National Tropical Botanical Garden,
United States
Reviewed by:
Joana Magos Brehm,
University of Birmingham,
United Kingdom
Nyree Zerega,
Northwestern University,
United States
*Correspondence:
Tinde Van Andel
tinde.vanandel@naturalis.nl
Specialty section:
This article was submitted to Plant
Systematics and Evolution,
a section of the journal
Frontiers in Plant Science
Received: 27 February 2019
Accepted: 26 August 2019
Published: 20 September 2019
Citation:
Van Andel T, Veltman MA,
Bertin A, Maat H, Polime T,
Hille Ris Lambers D, Tjoe Awie J,
De Boer H and Manzanilla V (2019)
Hidden Rice Diversity in the Guianas.
Front. Plant Sci. 10:1161.
doi: 10.3389/fpls.2019.01161
Traditional crop varieties are an important source of genetic diversity for crop adaptation
and modern breeding. Landraces of Asian (Oryza sativa) and African (Oryza glaberrima) rice
have been well studied on the continents where they were domesticated. However, their
history of cultivation in northern South America is poorly understood. Here, we reveal the
rice diversity that is maintained by Maroons, descendants of enslaved Africans who fled to
the interior forests of the Guianas ca. 300 years ago. We interviewed subsistence farmers
who practice shifting cultivation along the Maroni and Lawa rivers that form the natural
border between French Guiana and Suriname, and used ethnobotanical and morphological
methods to identify around 50 varieties, of which 15 were previously undocumented.
The genetic origin of these varieties was explored using the Angiosperms353 universal
probe set. Despite the large distances between sites and relative inaccessibility of the
area, phenotypic and genetic diversity did not display any geographic structure, which is
consistent with knowledge of seed exchange among members of the same ethnolinguistic
group. Although improved US cultivars were introduced in Maroon villages in the 1940s,
these have not displaced the traditional landraces, which are cherished for their taste and
nutritious qualities and for their importance in Maroon spiritual life. The unique agricultural
and ritual practices of Maroons confirm their role as custodians of rice diversity, a role
that is currently under threat from external pressures and encroaching globalization. We
expect that the rice diversity uncovered in this study represents only a fraction of the total
diversity in the Guianas and may constitute a large untapped resource that holds promise
for future rice improvement. Further efforts to inventory and preserve these landraces will
help to protect a precious cultural heritage and local food security.
Keywords: ethnobotany, French Guiana, landraces, Maroons, PAFTOL, rice, Suriname, traditional agriculture
INTRODUCTION
Rice is the most widely consumed staple food in the world. Two species of domesticated rice exist: the
widely cultivated Asian rice (Oryza sativa L.), domesticated in China some 10,000 years ago, and the
lesser-known African rice (O. glaberrima Steud), domesticated about 3,000 years ago (Stein et al., 2018).
There are thousands of rice cultivars in Asia, Africa, and the Americas, and a significant proportion
of this diversity is maintained in germplasm institutes (Jackson and Lettington, 2002; Sanchez et al.,
2013). With a growing world population and increasing impacts of climate change, rice breeders
urgently need to develop more sustainable cultivars with higher yields, healthier grains and reduced
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Commercial rice cultivation in the Guianas started only in
the early 1900s in the coastal swamps of Guyana and Suriname.
The first improved cultivars were based on traditional varieties
brought by indentured laborers from India and Java, who were
recruited to work on the plantations after slavery was abolished
(Codd and Peterkin, 1933; Stahel, 1933). From the 1950s onwards,
commercial Asian wetland cultivars suitable for mechanical
harvesting were planted in newly constructed polders in
Suriname (Young and Angier, 2010; Maat and Van Andel, 2018).
Commercial rice fields in French Guiana were not developed
until 1982, with the construction of coastal polders around
Mana (Clément et al., 2011). Today, new cultivars continue to be
developed by the Anne van Dijk Rice Research Centre (SNRI/
ADRON, www.snri-adron.com), the national institute for rice
research in Nieuw Nickerie, Suriname.
Traditional Maroon agriculture is under pressure by increasing
commercial and governmental interventions. Gold mining and
logging concessions have been issued on traditional territories,
while prospects of education and employment stimulate
migration to urban areas (Paramaribo, Cayenne, St. Laurent),
which in turn leads to a shortage of farm labor (Heemskerk,
2000; Fleskens and Jorritsma, 2010; Price, 2012). Recent
infrastructural developments (roads, airstrips) have facilitated
access to the remote hinterland (Price, 2012) and commercially
produced rice is now widely available in gold miner shops in
the interior (Heemskerk, 2000). Shortening of fallow periods
in shifting cultivation plots and limited use of fertilizers keep
soil productivity low, with yields rarely exceeding 1000 kg/ha
and arguably not meeting local demand (Baumgart et al., 1998;
Fleskens and Jorritsma, 2010; Nascente and Kromocardi, 2017).
The increasing influence of evangelical churches in Maroon
territory, development organizations and government policies
have resulted in the perception of traditional Maroon farmers
as ‘backward’ and in need of modernization (Heemskerk, 2003;
Fleskens and Jorritsma, 2010; Léobal, 2016). These factors
actively discourage traditional Maroon farming, and put their
landraces at risk of disappearing.
No systematic inventory of Maroon rice landraces with storage
of vouchers or germplasm has been made thus far and their
varieties have not been included in formal breeding experiments
or field trials. In 1936, a French agronomist discovered more
than 30 rice varieties cultivated by Maroons along the Maroni
and Tapanahoni rivers (Vaillant, 1948). Almost 20 years later,
Portères (1955) identified these collections as Oryza sativa, except
for one sample that represented African rice (O. glaberrima).
Vaillant (1948) made an urgent appeal to continue the study of
Maroon rice varieties, as they might ‘represent a detached branch
of historical African cultivars at a time where European culture
and importation of selected hybrids had not yet played a role’,
but this was not followed up. Anthropologists Hurault (1965),
Bilby et al. (1989), and Price (1993) mentioned Maroon rice
diversity, but hardly published any landrace names. Biologist
Geijskes (1955) recorded 23 names of rice varieties along the
Maroni River, and anthropologist Fleury (2016) listed 21 names
among the Aluku on the Lawa River, but no herbarium or seed
collections were made. The only expedition to collect Maroon
rice varieties was carried out by Baumgart et al. (1998) along the
environmental footprints (Stein et al., 2018; Wang et al., 2018).
The narrow genetic base of modern crop cultivars guarantees
phenotypic uniformity and genetic stability, but also makes them
vulnerable to environmental fluctuations, such as climate change,
reduced soil fertility, pests, and diseases (Zeven, 1998).
Wild relatives of rice and traditional landraces often show
adaptations towards marginal environments and pest resistance
and are therefore considered as an untapped genetic resource for
breeding new cultivars resilient to future challenges (Alvarez et al.,
2007; Wang et al., 2018). Landraces are also of key importance in local
food security and preserving cultural heritage (Perales et al., 2005;
Ardenghi et al., 2018), and reveal past migration patterns of humans
and their contacts with outsiders (Westengen et al., 2014). Over the
last few decades, a severe genetic erosion of crops has taken place due
to the replacement of landraces by modern cultivars (Zeven, 1998;
Dyer et al., 2014). Much effort has been made to safeguard landraces
ex situ (in gene banks) to make their genetic resources available
for breeders, but the in situ conservation of agrodiversity within
traditional farming systems has not been pursued to the same extent
(Maxted et al., 2002). Ethnobotanical inventories are powerful tools
in detecting these neglected genetic resources and understanding the
social and cultural factors involved in generating and maintaining
their diversity and distribution (Westengen et al., 2014; Ardenghi
et al., 2018). Here, we describe the rice landraces that are grown by
Maroons, descendants of escaped slaves in Suriname and French
Guiana and discuss their efforts and motivations to maintain
this diversity.
Rice has been grown for centuries in the Guianas (Guyana,
Suriname, and French Guiana). In the 17th and 18th centuries,
plantation owners imported the crop from the US and West
Africa as provision for their enslaved laborers (Carney, 2009; Van
Andel et al., 2016a). Asian rice was introduced by Portuguese
sailors in West Africa in the 16th century, and both African and
Asian rice were grown by African farmers before the onset of
the transatlantic slave trade (Linares, 2002). Rice was sold in
the husk, allowing for longer storage and germination ability
(Carney, 2009). After their arrival in the Americas, slaves
managed to gather leftover seeds from ship’s hold or other
storage places and planted these in small provision grounds at the
periphery of the plantations (Price, 1991; Van Andel et al., 2012).
These home gardens enabled them to grow at least some of the
familiar crops of their motherland and became central to their
physical and spiritual life. The rice they cultivated consisted most
likely of upland, rainfed varieties (Carney, 2009; Fleury, 2012).
When they escaped from the plantations, Maroons brought crop
seeds and established new gardens in their communities along
the major rivers in the forested interior of Guyana, Suriname,
and the French side of the Maroni River (Codd and Peterkin,
1933; Hurault, 1965; Price, 1991). Today, there are no Maroons
left in Guyana, but six ethnic groups of Maroons still remain in
Suriname and French Guiana: Paramaccans, Aluku (or Boni),
Kwinti, Matawai, Ndyuka (or Aucans), and Saramaccans (Price,
2013). Rice has been their staple food for centuries, enabling
them to survive in the remote hinterland, independent from
coastal societies (Price, 1993; Fleury, 2012). In 2013, Maroons
numbered some 210,000 people and constituted around 25% of
the population of Suriname and French Guiana (Price, 2013).
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upper Suriname River, who deposited their samples at the SNRI/
ADRON seed bank. The discovery of a field of O. glaberrima in a
Saramaccan village in 2008 (Van Andel, 2010) stimulated further
ethnobotanical research on African crops grown among Maroons
(Van Andel et al., 2016b) and rice in particular (Van Andel et al.,
2016a; Maat and Van Andel, 2018), but genetic analysis of rice
was limited to one accession.
To complement these previous anthropological and
ethnobotanical studies, plants can now be studied through the
lens of genomics. Next generation sequencing analysis can reveal
further details about the origins, evolution, and diversity of
different crop varieties. Specifically, universal probes have recently
been designed for target enrichment of 353 low copy nuclear genes
across the Plant and Fungal Trees of Life (PAFTOL) (Johnson
et al., 2019). Target enrichment is a versatile, reproducible, and
scalable technique that increases the representation of selected
portions of the genome prior to its sequencing (Lemmon and
Lemmon, 2013). The Angiosperms353 probe set allows the
generation of high-quality genomic data for all the angiosperm
taxa, making results comparable at an unprecedented taxonomic
breadth, but its utility for differentiating closely related samples
of the same species has not yet been demonstrated.
In this study, our first aim was to collect herbarium vouchers,
seed material, and DNA samples of different rice species and
landraces grown by Maroons along the French Guiana-Suriname
border. Our second aim was to document their morphological
characters and associated traditional knowledge on their agricultural
requirements, cultivation and processing methods, local names and
meanings, culinary and spiritual values. Based on previous studies,
we anticipated that Maroon rice varieties would be known under
a range of local names and hard to tell apart visually. We therefore
tested the ability of the Angiosperms353 markers to capture the
genetic diversity of these traditional landraces and compared these
with other commercial cultivars and traditional varieties grown in
the region, demonstrating the functionality of the Angiosperms353
universal probe set (Johnson et al., 2019) for population genomics.
Local rice farmers were recruited by our translators, who
explained our research objectives in the Ndyuka or Saramaccan
language if they were not fluent in Dutch, French, or Sranantongo
(the lingua franca in Suriname). Having obtained the farmers’
oral prior informed consent, we conducted face-to-face
interviews and visited their rice fields if they had a standing crop.
Herbarium vouchers of living rice plants were collected using
standard botanical methods, and one duplicate was deposited at
the Herbier IRD du Guyane (CAY) in Cayenne, French Guyana
and the other in the herbarium of Naturalis Biodiversity Center
(L) in Leiden, the Netherlands. When no living plants were
available for specific varieties, we collected seeds from rice
stored in people’s outdoor kitchens. Seed samples were stored
in paper envelopes: one duplicate of each variety was sent to
the SNRI/ADRON germplasm bank in Suriname for storage
and phenotyping, while the other was stored at the Economic
Botany collection of L. We used the ‘rice passport data table’,
developed by the SNRI/ADRON, to document local names and
agronomical and culinary characteristics of each collected rice
variety. We expanded this table into a semi-structured interview
by adding questions concerning positive and negative features of
landraces (Supplementary Table S1). We also posed additional
questions about challenges faced in rice cultivation, the use of
agrochemicals, and the importance of rice in ancestor rituals.
For each collected rice variety, we documented morphological
characteristics in the field (size, husk and pericarp color,
presence of awns, anthocyanins, etc.) and made detailed
pictures of living plants, panicles, and husked and dehusked
seeds. To facilitate the discussion in the field, we made a ‘rice
book’: a ring binder with samples of local rice varieties secured
under transparent tape (Supplementary Figure S1). We made
short videos on the different stages of rice cultivation and
processing and uploaded these on YouTube. To raise awareness
on local rice diversity, we designed a poster with pictures of the
different rice varieties and associated traditional information,
which will be distributed in Suriname and French Guiana
(Supplementary Figure S2). Throughout this paper, we follow
Zeven (1998) and use the term ‘landrace’ for farmer-developed
rice accessions, ‘cultivar’ for rice accessions developed by
companies or breeding institutes, and ‘varieties’ for all rice
accessions, regardless of genetic improvement.
MATERIALS AND METHODS
Ethnobotanical Survey
Before the start of the fieldwork, a prior informed consent
form was prepared in French following the guidelines of the
Collectivité Territoriale de Guyane in Cayenne. This document
was signed on 10 April 2017 by Mr. Jacques Chapel Martin,
captain of the village of Grand Santi and responsible for the
Ndyuka traditional authorities of the region Grand Santi (Lawa
River). St. Laurent du Maroni is considered an urban area where
no traditional authorities are in place (Tareau et al., 2017), so
no such document was needed. Fieldwork was done from 4 to
22 July 2017. We interviewed Maroon rice farmers around St.
Laurent and Bigiston (lower Maroni River), near Providence
(upper Maroni River), near the Gonini River mouth, and around
Grand Santi (Lawa River), located ca. 144 km upstream from
St. Laurent (Figure 1). These locations were chosen because
of their relatively easy access and previous contacts with local
Maroon farmers.
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Genetic Analysis
Fresh leaf material from living rice plants was stored on silica
gel. In five cases, we germinated seeds on wet tissue paper and
extracted DNA from the green sprouts. DNA was extracted from
approximately 40 mg of dry leaf material using the DNeasy Plant
Mini Kit (Qiagen). Total DNA (0.2–1.0 µg) was sheared to 500 bp
fragments using a Covaris S220 sonicator (Woburn, MA, USA).
Dual indexed libraries were prepared using the protocol of Meyer
and Kircher (2010) for shotgun sequencing and target capture.
We captured target sequences with the PAFTOL project probe
set (Johnson et al., 2019). We prepared and pooled 41 equimolar
libraries in two capture reactions with an average 300 ng of input
DNA per pool. The RNA probes were hybridized for 24 hours
before target baiting, and 14 PCR cycles were carried out after
enrichment following the MyBaits v.3 manual. The enriched
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FIGURE 1 | Study area. Fieldwork locations are represented with green circles, whose size is proportional to the number of samples. The pie charts illustrate the
proportion of samples with different husk colors and anthocyanins. Metadata for this map are provided in Supplementary Table S3.
libraries were sequenced on one Illumina HiSeq 3000 lane
(150bp paired-end).
To find out whether the Maroon rice varieties bear genetic
similarity to other Asian rice grown in our study area, we
included 10 accessions from French Guiana and Suriname for
which genetic data is freely available (Supplementary Table S2).
These accessions were resequenced as part of the 3000 Rice
Genomes Project (3KRGP) and represent a mix of populations
and subspecies, including Oryza sativa ssp. japonica (4), O. sativa
ssp. indica (4), and two admixed individuals between O. sativa
ssp. japonica and O. sativa ssp. indica (Wang et al., 2018). The
raw sequencing reads were trimmed and quality filtered using
Trimmomatic v.0.32 (Bolger et al., 2014) and mapped with the
Burrows-Wheeler Aligner v.0.7.5a BWA-MEM algorithm (Li
and Durbin, 2009) against the reference genome of Oryza sativa
subsp. indica (ref. ASM465v1). The BAM files were filtered to a
minimum quality score of 30 and read depth of 5 with samtools
v.1.3.1 (Li et al., 2009). SNPs were called with ANGSD v.0.549
(Korneliussen et al., 2014) and filtered with vcftools v.0.1.13
(Danecek et al., 2011). Sites with more than 20% missing data
were discarded. The parameters used for the SNP filtering were:
minimum depth: 5, maximum depth: 500, minimum mapping
quality: 30, minimum minor allele frequency: 0.1. SNPs were
pruned with PLINK v.1.90b5.2 (Purcell et al., 2007) using a
pairwise correlation threshold of 0.5 in sliding windows of 500
SNPs with step size 50.
Multi-dimensional scaling of the genetic variation was
performed in PLINK v.1.90b5.2. Population structure analyses
were conducted with ADMIXTURE v.1.3 (Alexander and
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Lange, 2011). Genetic differentiation between sampling locations
was estimated with Weir and Cockerham’s weighted Fst statistics
(Weir and Cockerham, 1984) as implemented in vcftools v.0.1.13.
The average genetic diversity per population was calculated as π
in vcftools and tested for significance with the non-parametric
Kruskal-Wallis test. To ensure independent measures, we chose a
window size of 150 kb, which exceeds the average extent of linkage
disequilibrium as estimated in O. sativa ssp. japonica (Mather et al.,
2007). An maximum-likelihood (ML) tree was constructed in
FastTree v.2.1 (Price et al., 2010), after removing heterozygous sites
with bcftools v.1.1 (Li et al., 2009) and concatenating SNPs with a
custom perls script (Bergey, 2012), and using the color scale cividis
(Nuñez et al., 2018). Pairwise genetic distances were computed
using the p-distance model in MEGA7 (Kumar et al., 2016).
All scripts and command line options used for the analyses are
openly available on Zenodo (doi: 10.5281/zenodo.3276199). Raw
sequence reads are publicly available at: http://www.ncbi.nlm.nih.
gov/bioproject/554407 (Bioproject ID: PRJNA554407).
RESULTS
Farming Practices
We interviewed 21 rice farmers, 20 of Ndyuka and 1 of Saramaccan
descent, of which 8 were in the lower Maroni region (St. Laurent,
Bigiston, Manjabon), 3 near Providence, 6 in Grand Santi, and 4
near the Gonini River mouth (Figure 1). All participants were
female except one, who showed us the field of his sister-in-law.
No male rice farmers were mentioned, as it is the Maroon women
who do the planting, weeding, harvesting, and processing of rice,
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either by themselves or together with female relatives. Men do
help with the clearing and burning of fields in the dry period
around November. Rice is sown as the first crop in the fertile
ashes at the onset of the short rainy season between December
and January. After weeds and debris have been removed by
hand, rice is directly sown in the soil, either in small holes
(‘diki olo’) dug with a hoe (‘tyap’), or cast (‘fringi’) over a tilled
field (Video 1, see section 10). Rice grains must be covered by
loose soil and pressed flat to protect them from birds that feed
on exposed seeds. One farmer assured us that birds in primary
forest are not yet accustomed to rice fields, making it possible
to sow the grains without covering them. Most rice is sown in
well-drained, sandy upland soils on fields cut in primary forest
or in secondary forest in various stages of succession. Three of
the 21 interviewed farmers cultivated rice on brown clay soils in
a floodplain forest, which was very successful, while one farmer
grew her crop in a swampy white sand savanna, with meager
results. In Grand Santi (French Guiana), some women said they
do not go to their rice fields on Fridays, as this is prohibited by the
‘local God of the forest’. Therefore, some people also have a rice
field on the Surinamese side of the Lawa River, as there the bush
spirits forbid entry on Sundays. This way, they can work on their
rice fields every day of the week. Given the numerous islands,
unclear borders, and absence of border control, many Maroons
(temporarily) live and practice agriculture on both sides of the
Lawa and upper Maroni rivers.
Generally, rice is harvested between March and May. In St.
Laurent, other fields are then directly cut and burned, so a second
rice crop can be harvested from July to early September. After
this, no more rice is planted until December. In older fields, the
rice grains that shatter during the first harvest from March to May
will sprout between the cassava plants and mature four months
later. Because of the lack of nutrients, this ‘grow-back rice’ has
meager-looking stems with few-seeded panicles. The crop is also
much shorter (80–90 cm) than rice sown on freshly burned fields
(>1.60 m), but harvested nevertheless. Farmers in Grand Santi
do not plant a second rice field after their first harvest in April, as
there is too much rain in July for the rice to mature. Precipitation
data (World Bank Group, 2018) indeed show a slightly higher
rainfall in July in Grand Santi (198 mm) than in St. Laurent (186
mm). Since no mature rice fields were present around Grand
Santi during our fieldwork in July, we made our herbarium
collections in this region from ‘grow-back rice’.
Farmers sow their varieties one after another on different
parts of their fields and harvest them sequentially to divide the
workload over time and facilitate separate storage. None of the 21
farmers we interviewed used any pesticides or fertilizers. While
most varieties mature within 4 months, the white Alekisola,
Watralanti, and Mesti take longer, so they are often sown first
and harvested last. The ripe ears are cut off with a small knife
(Figure 2, Video 2), tied to sheaves (Figure 3), and dried
for some time in the sun on corrugated iron sheets or woven
polypropylene rice bags. When fully dry, the bundles are stored
separately per variety in plastic buckets, steel oil drums, glass
bottles, or rice bags. According to the farmers, some varieties
take longer to dry than others. Freshly picked rice is sometimes
consumed the same day, but then the ears are dipped in boiling
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FIGURE 2 | Harvesting ripe panicles by hand near Bigiston, lower Maroni
River. Picture by Alice Bertin. Written informed consent was obtained for the
publication of this image.
FIGURE 3 | Dried rice tied into sheaves, St. Laurent. In the background a
plastic drum in which dried rice is stored for next year’s planting stock and
consumption. Picture by Alice Bertin. Written informed consent was obtained
for the publication of this image.
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taste, as the machine mill removes all the bran, ‘and that is the
healthy part of the rice’. They still prefer mechanically milled
home-grown rice over shop rice.
Locally grown rice has symbolic value too. Funerals are
important social events among the Maroons and may last several
days, so relatives from faraway are able to attend. When someone
dies in a traditional Maroon village, the family members are
required to bring home-grown rice to the mourning ceremony
(Price, 1993). A gift of shop-bought rice or machine-milled
traditional rice is not accepted. Before the burial takes place,
women jointly pound and winnow the rice and prepare huge
quantities of different rice dishes, a part of which is placed on a
banana leaf as a sacrifice for the deceased, while the remainder
is consumed (Fleury, 2012; Reijers, 2014). Attendance of funerals
was frequently mentioned as a motivation for growing rice, even
by those who identified themselves as ‘church people’.
All but three of the farmers we interviewed obtained their
planting stock as a gift from family members. From the other
three, one bought it in a neighboring village and two purchased
it on the Charbonnière market. Most Maroons in St. Laurent and
Grand Santi are migrants who obtained their rice varieties from
relatives in traditional villages deep in the interior of Suriname, as
far as Tyontyon island and Mpuusu (Ndyuka territory, Tapanahoni
River) and Masia Creek (Saramaccan territory, upper Suriname
River). One variety (Aluku paansu) was exchanged with Aluku
Maroons on the upper Lawa River. None of the farmers reported
having bought seeds in Paramaribo or Cayenne.
water to facilitate removal of the husks. Farmers said that when
still in their tough, fibrous husk, rice seeds can be stored for
as many as 3 years without spoiling or losing their ability to
germinate. Sowing stock (‘paansu’) is not stored separately from
the rice meant for consumption.
Threshing is done before storage (when seeds are stored
separately in oil drums) or when it is time to consume the
rice. Bundles are spread out on a plastic sheet and the seeds
are separated from the ears by spreading out the sheaves and
trampling on them barefoot with a dancing movement (Video 3).
Rice varieties with scabrous stems are sometimes threshed by
putting the bundles in a bag and beating it with a stick. Only just
before consumption, the rice is hand-dehusked by pounding it
with a pestle in a heavy wooden mortar (Video 4). To winnow
the rice, it is put on a large wooden tray and thrown into the
air to let the wind blow away the empty husks (Video 5). The
pounding process can be repeated to also remove the bran
(pericarp), and get ‘clean and very white rice’, with broken grains,
if this is preferred. Some villages have diesel-powered rice mills,
for which the farmers have to pay € 4 to mill a 20-liter tin of rice.
Motivations for Growing Rice
A major motivation for growing rice is that it is much cheaper
than buying it in a store. In most Maroon families, rice is eaten
with every meal. The farmers we interviewed said that with a
good harvest, they would have sufficient rice to last for an entire
year or to the next harvest. Some women kept more than two full
oil drums (200 liters each) of rice grains in stock in their outdoor
kitchen. A few occasionally sell rice to other Maroons who do not
grow (enough) rice to be self-sufficient. On the ‘Ndyuka market’
on Charbonnière Plage in St. Laurent, 2-kg bags of milled Maroon
rice were sold for € 5 per bag. Most farmers said they only buy
rice when their own stocks are depleted. Since most shopkeepers
in the interior of Suriname and French Guiana are Chinese
entrepreneurs, shop rice is known as ‘Chinese rice’, although it
is commercially produced wetland rice from Nickerie (mostly
‘Breuk III’, a cheap type of broken rice of the Manglie brand). A
25-kg bag of this broken rice sells in the interior for € 20, which is
more than twice the price paid in Paramaribo (€ 7–8).
Almost all farmers we interviewed said that their home-grown
rice is healthier and tastier than the type that is sold by local
shops. People complained that ‘Chinese rice’ tasted ‘horrible’,
often had sand or stones in it, could not be stored long, and was
frequently infested by beetles or larvae. They also warned us that
‘eating only shop rice will give you disease’. Farmers repeatedly
assured us that Maroon rice ‘has vitamins and makes your belly
feel full’, has much ‘goma’ (starch), and ‘gives good milk’, referring
to the foam produced by the bran of home-grown rice during
cooking. One farmer remarked that the increasing number of
diabetes patients in St. Laurent should eat traditional rice instead
of shop rice to control their disease. Milling traditionally-grown
rice varieties by hand with a mortar and pestle removes the husk
but not the bran and the germ, so it is in fact ‘whole-grain rice’,
which is higher in fibers, lipids, proteins, minerals and vitamins
than polished rice and gives a fuller feeling after consumption
(Heinemann et al., 2005; Ryan, 2011). Farmers remarked that
when they mechanically process their rice, it loses some of its
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Challenges in Growing Rice
Many of the rice farmers we interviewed were young women
in their 20s or 30s, who said they enjoyed farming and were
dedicated to continue the practices that their ancestors had
passed on to them. They sometimes criticized others who
‘became too lazy’ to practice traditional agriculture or mill rice by
hand because of the French child support payments that allowed
them to buy rice in shops. None of the farmers complained about
shortages of farm labor or a lack of suitable areas to burn a field.
Birds are considered a minor nuisance and insect pests were
not reported. Although most interviewed farmers (14 of the 21)
had a rice field within walking distance from their home, 7 had
fields that were located further away and could only be reached
by boat. In spite of the high fuel costs, this was still considered
economically feasible. The only farmer who complained about
decreasing soil fertility had a field that was invaded by saplings
of Acacia mangium Willd. This Australian tree was introduced
to French Guiana in the 1970s to restore degraded mining areas
but became invasive. As fire activates its germination, it quickly
colonizes new agricultural fields (Delnatte and Meyer, 2012).
Locally known as ‘Mira udu’ (ant wood), it dominates secondary
forests in the outskirts of St. Laurent. Farmers said the only way
to get rid of it is by ring-barking. We did not observe A. mangium
elsewhere along the Maroni or Lawa River. A major challenge
mentioned by Maroon farmers around St. Laurent is harassment
by the French police. They said that during the dry season, the
authorities fly small airplanes over the area to detect burning
fields. Fines can be up to € 10,000, so some rice fields are made
far from home to avoid discovery.
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Diversity of Rice Varieties
We collected 38 herbarium vouchers of rice plants from 21
agricultural fields and 47 seed samples from living plants, stored
and dried panicles, or loose seeds. In Supplementary Table
S4, all collections are listed with their local name, collection
number, locality, and associated ethnobotanical and agronomical
information collected by means of the field questionnaire
(Supplementary Table S1). Based on morphological characters,
genetic analysis and local names, we grouped our collections
into ca. 50 varieties (Supplementary Table S4). From the seed
collections and herbarium vouchers with mature seeds, 56 seed
samples were sent to SNRI/ADRON. Of these, 48 samples were
successfully germinated by the SNRI/ADRON staff in their
research station in Nickerie (Suriname), grown into fertile adult
plants and phenotyped in July 2018. The number of different
rice varieties encountered per farmer (on the field and/or in
storage) ranged between 1 and 7, with a mean of 3.2. Several
farmers indicated that they had grown more varieties that year
(up to 9), but some stock had been depleted. Most of the varieties
were encountered only once. We extracted DNA from 40 rice
individuals of different local varieties (Supplementary Table S4).
Target sequences were captured from 35 herbarium vouchers and
five germinated seed samples. One individual was removed from
the analysis due to insufficient data. Following quality filtering,
the median sequencing depth was 5.5X, with a range of 1.1–9.8X,
resulting in a data set consisting of 39 individuals with 250,218
single nucleotide polymorphisms (SNPs).
FIGURE 4 | Oryza glaberrima seeds (nr. 6782), showing the black husks and
straight awns. Picture by Tinde van Andel
varieties analyzed, 37 form a highly supported monophyletic
clade with the 3KRG tropical japonica accessions (Figure 5A).
The short pairwise distances within this clade indicate relatively
recent divergence and stand in sharp contrast with the large
pairwise distances to the other 3KRGP admixed and indica
accessions (Supplementary Table S4). Principal component
analysis confirms that the collected upland rice is similar to, but
not identical with the tropical japonica accessions of the 3KRG
project (Figure 5B). This is also reflected in the population
structure analysis, which shows that the collected varieties share
a higher proportion of genetic ancestry with the tropical japonica
than with the indica and admixed accessions (Figure 5C). Only
one collected sample stands out: Watralanti (nr. 6754), which
means ‘water land’ and appears to be a wetland variety that bears
more similarity to the 3KRGP indica (Figure 5A Group I) and
particularly the admixed varieties Acorni and Ciwini (Figure 5A
Group II) that were developed in Suriname in the 1970s
(Supplementary Table S2). Watralanti is always planted near a
creek or at the bottom on a slope as it needs moist soil. This rice
probably descends from a wetland variety that was exchanged with
East Indian rice farmers in coastal Suriname several decades ago.
Since our study area was divided in six collection sites, we
checked for systematic differences in genetic diversity between sites.
There were no differences in pairwise genetic distances within
a geographic group (0.0137) as compared to between geographic
groups (0.0135), or to the population wide average (0.0136).
This shows that the variation of landraces is mixed evenly across
different sampling locations, and not structured geographically.
To test whether locations that are close together are more likely to
exchange rice genetic diversity, we assessed genetic differentiation
between populations along the North-South axis, following the
Lawa/Maroni watershed that serves as the main highway for
transportation of people and goods. In line with the low genetic
distances among the tropical japonica accessions, the fixation index
(Fst) was found to be extremely low between sites and frequently
less than 1% (Table 1). This is exceedingly so between the island
populations near Providence, Grand Santi and Mofina, which
Species and Subspecies Identification
All but one sample belong to the Asian rice species Oryza sativa.
The only exception is Baaka alisi (nrs. 6782 and 6773A), which
represents a variety of African rice (O. glaberrima). Although
people said African rice was not often cultivated, we encountered
three farmers who kept small bags or glass bottles in stock with
this conspicuously straight-awned rice with black husks and red
pericarp (Figure 4). The use of African rice was shrouded in
some mystery. Two farmers said that ‘the ancestors came with
this rice, they brought it and ate it’. One farmer said that if you
pound the black rice well enough, it becomes white (as it loses
its red bran), so she would cook and eat it mixed it with her
homegrown Asian rice. Other people ensured us that black rice
is never eaten nor brought as a gift or prepared during funerals.
The rice is mostly sold for a good price (± € 1 for a small bag
of 40 gram on the Paramaribo market) to be used in traditional
medicine. People cook the milled rice into porridge and drink
the starchy water when they suffer from diarrhea. The husked
seeds are also used in herbal baths to get into contact with ‘deep
ancestor spirits’ during Afro-Surinamese rituals. Two of the three
farmers who grew black rice said they did not perform these
rituals themselves, because they were ‘church people’. Baumgart
et al. (1998) reported that black rice is only planted to keep the
birds away from the rest of the crop, but this seems unlikely, as it
is sold for a higher price than any of the other varieties.
The other varieties we collected were all identified as Asian
rice and, according to the farmers, mostly upland dry rice. Our
phylogenetic analyses demonstrate that these varieties probably
belong to the tropical japonica variety (Figure 5). Of the 38
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FIGURE 5 | Position of Maroon landraces within the Asian rice gene pool. (A) Approximately Maximum Likelihood (ML) tree of 39,595 homozygous SNPs. Branch
lengths are not shown. Branches with low support values (< 0.70) are collapsed. Support values of the remaining branches are indicated on a scale from 0.70
(red) through 0.85 (yellow) to 1.00 (green). (B) Principal component analysis of 64,313 unlinked SNPs. (C) ADMIXTURE analysis of 64,313 unlinked SNPs at K = 2
ancestral populations
TABLE 1 | Genetic differentiation between sampling locations. Fixation index was computed as Weir and Cockerham’s weighted Fst (Weir & Cockerham, 1984).
Sampling location
St. Laurent
Bigiston
Bigiston
Providence
Grand Santi
Mofina
Gonini
0.010833
0.0021599
0.02363
0.024592
0.061006
0.014761
−0.0034
0.012335
−0.00405
Providence
0.0068157
−0.0013556
0.030035
Mofina
−0.003736
−0.042314
0.006427
varieties beyond the subspecies divide (Supplementary Figure
S3B), but with the present data, this model is not supported.
When we zoom in on the genetic variation among the 37 rainfed
O. sativa collections, though, we do observe a clear separation
between a group of genetically and morphologically more
uniform improved varieties, that presumably entered Maroon
agriculture some 70 years ago (Figure 5A, Group III) and the
larger and more diverse group of traditional varieties (Figure
5A, Group IV). The internodal distance separating the improved
clade is noticeably larger than the internodal distances among the
traditional varieties (Supplementary Figure S4A). The narrow
genetic base of this group is particularly apparent when we limit
the two-dimensional scaling of genetic variation to the upland
rice varieties (Supplementary Figure S4B). In this analysis, we
see that most of the genetic space is occupied by the traditional
landraces, whereas the improved varieties cluster relatively close
together. To confirm that these modern varieties (Group III) are
genetically less diverse than the traditional landraces (Group
IV), we compared their average nucleotide diversity in 150 kb
windows to an equal number of randomly chosen individuals
exhibit almost complete panmixia. Population differentiation is the
highest (> 2%) between the far northern areas of St. Laurent and
the southern areas around Grand Santi and the Gonini river. This is
consistent with a view of the Lawa/Maroni river as a transmission
vector, along which rice is exchanged and diversity is actively
maintained. Rather than a barrier against gene flow, the river is
thus a place where gene pools meet and mix. This explains why the
genetic differentiation between the opposite sides and countries is
negligible, and why the legal border between French Guiana and
Suriname does not form a natural border that leads to isolation.
Improved Varieties
It is noteworthy that our collection of landraces (with the exception
of Watralanti), although morphologically heterogeneous,
appears to form a single large population without much genetic
or geographic substructure. The cross-validation error estimate
in ADMIXTURE was lowest at K = 2 ancestral populations,
with one of the ancestral populations accounting for most of the
genetic makeup of the varieties (Supplementary Figure S3A).
Only at K = 4, we begin to differentiate groups within the Maroon
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Grand Santi
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Maroon Rice Diversity
from the other group, using a one-tailed Wilcoxon Rank Sum
test. This showed that at 0.00011, the average divergence per
window of Group III is significantly lower than that of Group IV
at 0.00013 (p < 0.02) (Supplementary Table S5).
This is of interest, because Group III includes the two most
frequently cultivated varieties. The most popular one is known
as ‘Mesti’ and grown by 29% of the farmers we interviewed. This
term means ‘teacher’ in Ndyuka, but only one farmer remembered
that this variety was named after a teacher who came to work at
the first boarding school established in the region by the Moravian
church on Stoelmanseiland, the island located in the confluence
of the Tapanahoni and Lawa Rivers (Figure 1). As children from
remote Maroon villages stayed on the island during the semester
and there were few shops in the area, the teacher handed out rice to
the mothers to grow on their own fields. The rice was subsequently
supplied to feed the boarding school pupils. The white-husked Mesti
rice quickly became popular and is still grown today. Farmers said
it took longer to mature than most other varieties and compared
its thin, elongated, slightly curved grains to the commercial ‘semisuper’ rice sold in shops. We could not trace the name of the teacher
who introduced this variety, but it must have happened shortly
after World War II, when the Moravians started their first mission
center on Stoelmanseiland (Jabini, 2012). Vaillant (1948) did not
encounter this variety in 1936, but a Maroon rice variety named
‘Mesiti’ was reported in the area in 1952 (Geijskes, 1955). We do not
know the exact origin of Mesti rice.
Another popular and closely related variety is the ‘white
Alekisoola’ or ‘white Sola’, grown by four farmers. It is considered
as a ‘beautiful rice’, which is often sown earlier than other varieties,
as it takes long to mature. It is planted on slopes and cannot
tolerate drought. The rice does not break during dehusking
and its grains are said to be ‘long like the commercial rice from
Nickerie’. A drawback is that heavy rains during the harvest time
cause the grains to become black and brittle. Varieties with this
name were also reported by Geijskes (1955) and Baumgart et al.
(1998) and identified as locally adapted farmers’ selections of
Rexoro. This glabrous-hulled cultivar was developed in 1926 in
Louisiana (Rutger and Mackill, 2001), introduced to Guyana in
1932 (Codd and Peterkin, 1933), and widely grown in coastal
Suriname by 1938. According to Stahel (1944), a bale of ‘Rexora
seed padi’ was sent to the Saramaccan village Ganzee in 1936 and
a few years later it was already grown by Ndyuka’s in the Maroni
area. When Vaillant did his survey in 1936, it was not yet present
there. The cultivation of Rexoro was discontinued in the 1960s
because of a serious attack of Cercospora disease (Sanderson,
1962), but Alvarez et al. (2007) recently encountered it again
on traditional farms in Cuba, decades after the introduction of
modern varieties by the Cuban rice breeding program. The redhusked types of (Aleki-) Sola that we encountered have different
morphological features than the white-husked types. They are
sown and harvested together with the other rice varieties, with
which they are genetically more closely related (Figure 5A).
types of a specific traditional variety, which are sown, harvested,
and stored separately: a ‘white’ one with straw-colored husks and
a ‘red’ one, with orange-brown husks (Supplementary Table S4).
Some women assured us that the only difference between these two
types is the color, but our phylogenetic analysis (Figure 5) shows
that in no less than six varieties, the white-husked types (of Onini,
Sola, Alekisola, Alulu, Apiikutufutu, Abadagai and Bau anu) are not
nearest neighbors with the red-husked types, but end up in different
clades. In other cases, where varieties had the same name and the
same husk color (such as Alulu, Weti alisi and Lebi Alisi), we found
that they were phylogenetically separated. This shows that shared
local names are not necessarily rooted in shared ancestry.
Several varieties with purple stems are named ‘Bau anu’ (blue
hand), as the anthocyanins in the panicles stain people’s hands
blue during the harvest. These varieties are not related and cluster
phylogenetically with other varieties known under another local
name. The blue sap is not seen as a negative trait; blue hand rice is
said to be tasty, although it does not always give a good yield. One
of the blue hand varieties (no DNA sequenced) is also known
as ‘Ayengena’ (‘it does not hang’ in Ndyuka, after its somewhat
erect panicles). In the early 1900s, Codd and Peterkin (1933)
reported several varieties with purple pigmentation in Guyana,
but we lack information on whether these are related to presentday Maroon varieties.
Six morphologically different varieties are named ‘Alulu (a
bon)’, which literally means ‘it rolls (from the tree)’ or ‘it crumbles’
(Figures 6A, B, Supplementary Table S4). The ripe seeds of this
rice drop easily from the ear, so farmers cut the panicles just before
they are fully mature and immediately place them in a bag to prevent
the seeds from falling on the ground. The loss of seed shattering
was a key event in the domestication of rice, as shattering caused
a severe reduction of yield (Konishi et al., 2006). Maroon farmers,
however, see the shattering as a positive trait, as it facilitates the
threshing process. They praise this type of rice for its tasty, fat, white
grains and ease of milling, although it is not tolerant to drought.
The three ‘Alulu’ types that we sequenced cluster in different clades,
so the shattering trait may have been reacquired several times
independently or persisted from an ancestral gene pool.
Wintawaai (nr. 6809) is a variety that quickly lodges when
there is much wind during the harvest season, after which the
seeds shatter and rot. Lodging is generally seen as an undesirable
trait of inferior landraces (Kashiwagi et al., 2005), but according
to the single farmer who planted this variety, it grows well, is tasty
and just needs to be cut before it is fully ripe.
One type of red-husked rice (Lebi alisi, nr. 6800) found near
Providence, upper Maroni River, grouped together with the
historic Wanica variety that was sequenced by Wang et al. (2018)
in the 3000 Rice Genomes Project (Figure 5). This 4-month variety
was reportedly grown by Maroons in the 1930s in the interior of
Suriname (Stahel, 1933). Two of our Maroon varieties (Aluku
paansu and Abadagai) grouped close to one of the reference
varieties from French Guiana (IRIS_313_7922, Figure 5), but no
further information was provided on the origin of this accession.
Traditional Varieties
Ancestor Varieties
Whereas the traditional Maroon varieties are cultivated less
frequently, this is a larger group with higher genetic and phenotypic
diversity (Figure 5A, Group IV). Farmers often distinguish two
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In 1936, Maroons told Vaillant (1948) that according to their
legend, rice came from Africa and was brought to Suriname
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joined by a female runaway named Ma Sapá, who had hidden
rice kernels in her hair. The current Sapali rice likely descends
from an early 18th century variety that has been crucial to the
survival of the Ndyuka Maroons.
Obsolete and Weedy Rice Varieties
The exact number of Maroon rice varieties is probably higher
than 50, as we missed some varieties that had been planted in
2017, but had already been consumed, such as the red and white
types of the Saramaccan varieties Tjë́kë muyee and Agbonso.
Farmers assured us that many more varieties could be found
in traditional villages along the remote Tapanahoni and upper
Suriname Rivers. Some women told us about varieties planted
by their mothers that are not sown anymore nowadays, because
of their long, itching awns or scabrous leaves, which are seen as
a nuisance. Examples are ‘Saapa’ (probably referring to the Sapali
variety) and rice types aptly described as ‘pubic hair’ after their
long, curled black awns. The latter varieties (nrs. 6758, 6767)
were still grown by two of the 21 interviewed farmers, who said
that ‘they look beautiful, give a good yield and we enjoy planting
it’. One farmer said she was ‘surprised that people still grow
this old-fashioned rice’, when we showed her our collections
of awned, shattering and lodging varieties. Two of the tropical
japonica varieties that were sequenced by Wang et al. (2018)
go by a similar name (Pinde gogowierie). Like Wanica, many
of the 3KRGP accessions were collected and deposited in seed
banks several decades ago, which means they do not necessarily
represent the currently used gene stock.
In addition, there was much morphological variation in the
types that carried the generic names ‘weti alisi’ and ‘lebi alisi’
(white resp. red rice), but we were not able to extract DNA from
every single accession. On the upper Maroni islands near the
village of Providence (Figure 1), we collected two samples of
‘white rice’ (nrs. 6799, 6801) that turned out to be mixtures of
grains with pubescent and glabrous husks, dark and uncolored
tips, white and red pericarp (Supplementary Table S4). The redseeded rice (nrs. 6801A and 6799B) is sometimes called ‘lebi wata’,
referring to the red color of the water when it is cooked, but it is not
stored separately from varieties with white pericarp. Apparently,
the samples were infested with ‘weedy’ or ‘red’ rice, sometimes
indicated as Oryza sativa f. spontanea Roshev., although this name
is considered to be a synonym of O. sativa (The Plant List, 2013).
In the 1920s, the ‘predominance of a particularly undesirable
type of wild rice with red grains’ among traditional landraces was
one of the reasons for the selection of pure lines for commercial
rice cultivation in Guyana (Codd and Peterkin, 1933). Weedy
rice is still a serious agricultural problem in directly sown rice
worldwide, and various types exist (with or without awns and
with different husk colors), often mimicking the phenotype of the
desired variety (Londo and Schaal, 2007). These red-seeded types
can arise when Asian rice hybridizes with its wild ancestor, Oryza
rufipogon Griff., creating offspring that shatter quickly, have long,
barbed awns, increased competitive ability, dormant seeds and a
red pericarp. Weedy rice also develops when domesticated rice
is abandoned and turns feral (Ellstrand et al., 2003). The weedy
rice types we encountered probably belong to the ‘strawhull type’
(Gealy, 2005; Londo and Schaal, 2007), as they lack awns, have
FIGURE 6 | Two shattering varieties of Maroon rice named ‘Alulu’ with
husked (left) and dehusked (right) grains. (A) ‘White type’ with strawcoloured, pubescent, black-tipped husks (nr. 6772A) and (B) ‘Red type’,
with orange-brown, glabrous, black-tipped husks (nr. 6819). Pictures by
Alice Bertin.
by a female ancestor who concealed the grains in her hair
(Carney, 2009). We found this legend still to be known in St.
Laurent today, and videotaped a demonstration of how a Maroon
woman braids hands full of rice grains invisibly into her daughter’s
hair (video 6). She explained that besides rice, other crop seeds
and cassava cuttings were also smuggled this way, both in Africa
when women were enslaved and in Suriname when Maroons fled
to into the forest. Moreover, we encountered two rice varieties in
Bigiston that still carry the names of the enslaved women who
brought this rice to the interior. The variety ‘Milly’ (nr. 6760) is
named after an escaped female ancestor ‘who introduced this
rice to the Maroni River in times of slavery when people did
not yet live in the forest [ … ] when they had not yet created
villages in the woods’. Another variety, growing in the same
field, is named ‘Sapali’ (nr. 6761), after an enslaved woman who,
after she fled from the plantation, crossed a savanna where she
encountered rice plants, from which she took ripe seeds that she
shared with other runaways. Although Sapali is appreciated for
its ‘fat, white and clean’ grains, the scabrous leaves scratch your
skin and the seeds are difficult to dehusk by hand. According to
Ndyukas along the Cottica River interviewed around 1961 by
anthropologist Köbben (1968), their ancestors had suffered from
hunger after their escape from the plantations, until they were
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straw-colored husks and red pericarp. Although of Asian origin
(Stein et al., 2018), O. rufipogon occurs in the wild in the Guianas
(Judziewicz, 1990). The herbarium label of a specimen from 1978
(P. Grenand nr. 1633, CAY) indicated that Palikur Indians in
French Guiana consumed this wild rice in the past, although the
awns were difficult to remove. Future genetic analyses can prove
whether the weedy rice we encountered represent hybrids between
traditional varieties and wild rice or escaped rice gone feral. The
preceding sections show a remarkable array of modern, improved,
traditional and ancient landraces, ranging from commonly used
to almost disused. Phenotypic diversity and vernacular names are
distributed rather homogeneously both in phylogenetic (Figure 5)
and in geographic space (Figure 1).
Exchange of seed material within ethno-linguistic groups
generally occurs at higher frequencies than between different
groups, as seed exchanges are based on cultural preferences
(Labeyrie et al., 2014). Ndyuka farmers hardly trade rice with
other Maroon groups, and they ensured us that ‘Saramaccans
had different types of rice’. This is in line with recent sorghum
studies showing that the spatial distribution of landrace names
and genetic diversity of traditional crops are significantly
correlated with ethnolinguistic partition (Westengen et al., 2014;
Labeyrie et al., 2014). We therefore think that the diversity of
landraces that we encountered represents only the tip of the
iceberg of hidden rice diversity in the Guianas, and that similar
ethnobotanical surveys among other Maroon groups will reveal
additional varieties not covered in this study. To date, no voucher
material exists for rice varieties of the Paramaccan Maroons who
live along the middle Maroni River or the Aluku Maroons along
the upper Lawa. Only a few collections have been made among
Saramaccan Maroons (Baumgart et al., 1998; Van Andel et al.,
2016b), but no research has been done on rice grown by the
Matawai and Kwinti Maroons in central Suriname. Apart from
a paper on rice cultivation by Hmong migrants from Laos in
French Guiana (Salaün, 1999), no studies or herbarium vouchers
exist of traditional rice varieties currently grown by descendants
of Asian migrants in the Guianas. This study is therefore the first
step towards obtaining a full picture of the landraces that are still
in cultivation by different ethnolinguistic groups in the Guianas.
DISCUSSION
More Hidden Rice Diversity
Since the 1936 expedition by Vaillant, our study is the first
inventory of Ndyuka Maroon rice that is supported by herbarium
vouchers, living seeds safeguarded at a germplasm institute and
genetic analysis. We rediscovered only seven of the ± 30 Ndyuka
rice names documented by Vaillant (1948), five of the 21 Aluku
names listed by Fleury (2016), four of the 25 Saramaccan names of
Baumgart et al. (1998) and five of the 23 Ndyuka names published
by Geijskes (1955). On the other hand, 15 of the local rice names
that we documented had never been reported before. We cannot
tell which of the previously documented varieties have been lost
in the past 80 years, as our inventory covered only a small part
of the Maroni and Lawa Rivers, rice names are subject to change,
and Vaillant’s herbarium vouchers are not digitally available.
Price and Price (2017) suggested a substantial decline in the
number of Maroon rice varieties over the past decades, but no
collections or name lists are available to prove this assumption.
Further morphological and genetic comparison of historic rice
collections (such as Vaillant’s vouchers at the Muséum National
d’Histoire Naturelle in Paris) with recent samples and additional
field collecting are needed to provide a more complete overview
of the extraordinary rice diversity in this area, its dynamics in
space and time and the survival of ancient landraces.
Along the Maroni-Lawa watershed, farmers cultivate a large
number of rice varieties, each with its own morphological and
agronomical characteristics. This phenotypic diversity, ranging
from ancient African landraces (O. glaberrima) to relatively
recent introductions (US cultivars of O. sativa subsp. japonica and
a Hindustani wetland variety) reflects the history of the human
migration in the region and their contacts with outsiders. This
diversity also reflects the multiple objectives of rice cultivation
for farming households: it is simultaneously a food production
strategy, a social and ritual asset. Foreign varieties introduced at
different times in the history of the region have contributed to
increase the phenotypic diversity. The fact that the phenotypic
diversity described in this paper is apparent at all collection sites
suggests a certain level of gene flow between the populations
along the river, serving as a pathway for seed exchange. This is
concordant with the lack of geographic and genetic structure
observed in our population genomic analyses (Figure 5A).
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Where Did the Maroon Rice Varieties
Come From?
Several rice varieties collected during this study possessed traits
that are characteristic to traditional landraces, such as tall stems,
awns, scabrous leaves, pubescent husks, shattering seeds, small
grains, and a proneness to lodging. Awns and rough hairs help to
protect seeds from predation by birds and mammals, and aid seed
dispersal by clinging to animal fur. However, awns and scabrous
plant parts also hinder mechanical seed processing and storage, so
modern rice cultivars have been selected to be glabrous, awnless
or short-awned (Rutger and Mackill, 2001; Hua et al., 2015).
Greater plant height makes rice more susceptible to lodging,
which is considered a negative trait for machine-harvested rice,
so modern cultivars have been selected for short and sturdy
stems (Nascente and Kromocardi, 2017). However, lodging does
not pose a problem if panicles are cut by hand (Kashiwagi et al.,
2005). The development of non-shattering cultivars has also
been a strong focus of rice breeding programs, although modern
indica cultivars are still more prone to exhibit seed shattering than
japonica cultivars (Konishi et al., 2006). Typically, all Maroon
accessions we sequenced are tropical japonicas, and several of
them shatter quickly, just like our sample of Oryza glaberrima.
In 1927, the national rice breeding program in Guyana
started to develop pure lines from locally grown rice landraces
by removing volunteer plants and selecting non-shattering,
awnless strains with a high yield, strong straw, uniform plant
height and white grains to facilitate mechanical threshing and
milling. These new cultivars were handed out to farmers to
ensure the ‘elimination of traditional varieties with undesirable
characteristics’ (Codd and Peterkin, 1933:2-3). In the 1940s,
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through World War II, the Dutch started investments in
commercial rice breeding in Suriname. They imported the pure
lines from Guyana and improved cultivars from Java, Indonesia
and the US for their field trials, but also tested out some varieties
that were probably of Maroon origin, like Wanica. After the war,
modern wetland cultivars were developed for mechanized rice
cultivation in the coastal polders (Maat and Van Andel, 2018).
It is likely that some of the varieties we collected descend from
these pre-1927 landraces, although the manifold origins of the
Maroon rice diversity are difficult to trace. Europeans introduced
the crop in the Guianas in the early colonial period, but the exact
routes and motivations remain unclear (Maat and Van Andel,
2018). Plantation holders imported hulled rice from Louisiana
and Carolina as provision for the slaves (Codd and Peterkin, 1933;
Rolander, 2008). Jewish planters that were expelled from Brazil
may have introduced rice to Suriname when they migrated there
around 1667 (Young and Angier, 2010). Slave traders bought stocks
of rice from merchants along the West African coast to feed their
captives during the Middle Passage (Carney, 2009; Mouser et al.,
2015; Van Andel et al., 2016a). Oryza sativa was introduced to West
Africa in the 16th century, before the onset of the transatlantic slave
trade, and adopted by peoples along the upper Guinea Coast who
had previous experience growing the local African species (Linares,
2002). Vaillant (1948) suggested that most of the Maroon rice
diversity had an African origin. However, Saramaccan rice farmers
interviewed by Price (1993) said that while in their youth (in the
1920s), there were only a handful rice varieties (black rice was one of
them), diversity increased substantially in the 1960s when Maroon
men took up wage labor in coastal Suriname and returned with
planting stock exchanged with East Indian and Javanese farmers.
Therefore, part of the Maroon rice diversity may have originated
from landraces introduced by Asian contract laborers.
Recently, the genome of a single accession of black rice from
Suriname was compared to 109 accessions of O. glaberrima
collected across West Africa. A strong similarity was established
with a landrace grown in western Ivory Coast (Van Andel et al.,
2016a), which shows how genomics can reveal unwritten migration
histories of crop varieties. Applying comparative phylogenomics
to historic rice collections from the Guianas in museums and gene
banks and currently cultivated Maroon landraces, and comparing
them with traditional varieties and cultivars from Africa, Asia,
and the US can shed a light on the geographic origins of Maroon
lineages and reveal patterns of crop migration and adaptation that
have remained hidden for centuries. Special attention should be
paid to varieties that Maroons consider to have been introduced
by enslaved ancestors, such as Milly and Sapali for the Ndyuka
and Paánza rice for the Saramaccans (Price, 2002), of which one
accession is stored at the SNRI/ADRON seed bank.
field. Early and late maturing types are sown in succession to
even out labor during harvest time. In this way, there is a higher
chance that at least one variety will thrive, as the weather may be
unpredictable, and wet and dry seasons do not always start and
end at the same time over the years. Cropping patterns that mimic
ecological complexity are most effective at soil conservation and
promoting sustainability, as long as rotation periods are long and
human population pressure is low (Kleinman et al., 1995).
Governments and agricultural development programs
generally promote continuous cropping systems over shifting
cultivation (Kleinman et al., 1995) and Suriname is no exception
to this phenomenon (Fleskens and Jorritsma, 2010). After
visiting the Maroon rice fields, Geijskes (1955) claimed that
yields were not always sufficient and ‘due to the wasteful land use,
the agriculture of the Bush negroes has become a matter which
urgently needs the attention of the government’. Tropical plant
breeders Budelman and Ketelaars (1974) advised that dryland
rice cultivation in the interior should be terminated, ‘as this is
a difficult crop for permanent cultivation on these grounds’ and
recommended oil palm plantations as a more suitable cash crop.
In 1992, the French government made violent attempts to force the
Maroon refugees from the Surinamese civil war to leave the area
and sprayed pesticides on their rice fields (Léobal, 2016). Maroon
farmers around St. Laurent, illegal immigrants and legal French
citizens alike, considered the ongoing harassment by the French
police as a serious problem affecting their agricultural practices.
As a remedy for short fallow periods and rice yields that did not
meet the local demand, caused by population pressure, Baumgart
et al. (1998) suggested to intensify Maroon rice cultivation by
purifying traditional varieties, introducing improved upland
cultivars, fertilizers, herbicides, and mechanization. However,
in their study on soil fertility on Maroon fields, Fleskens and
Jorritsma (2010) discovered that due to migration to the city, the
issue of human pressure on the land was rarely raised by Maroon
farmers. In a recent experiment, Nascente and Kromocardi (2017)
tried to increase yields by adding NPK fertilizer, herbicide, and
fungicide to three traditional Saramaccan landraces and three
improved Brazilian upland cultivars. Only one of the Maroon
varieties responded positively to this treatment, after which
they recommended to switch to Brazilian cultivars to double
the current average of upland rice production and improve food
security among the Maroon population. They acknowledged,
however, that the ‘very limited financial capital and low education
level’ of the Maroons limited their access to modern technologies
and that the introduction of high-yield rice cultivars required
subsidies for farmers to access fertilizers and pesticides (Nascente
and Kromocardi, 2017).
In addition to the governmental pressure, the upcoming evangelical
churches in the Maroon territories are rigidly opposed to Afroreligious practices, which has led to frictions within communities
(Van Stipriaan, 2015). According to Richard Price (pers. comm. 3
May 2018), converted Christians are increasingly discouraged to visit
funerals in remote Maroon villages, as these are regarded as places
of witchcraft. Since funerals are core events at which landraces are
exchanged, processed and consumed, this phenomenon could affect
the survival of Maroon rice diversity. Traditional crop varieties
can quickly disappear if not sown every few years. In Grand Santi,
Future of Maroon Rice
The Maroon forest garden represents the classical slash-and-burn
agroecosystem (Kleinman et al., 1995), in which rice is sown
once on a freshly burned field and intercropped with cassava,
maize, bananas and vegetables. Without the use of fertilizers or
pesticides, the field is left fallow after one season, after which only
volunteer rice plants and perennial crops are harvested. Farmers
spread their risk by planting several distinct rice varieties on one
Frontiers in Plant Science | www.frontiersin.org
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September 2019 | Volume 10 | Article 1161
Van Andel et al.
Maroon Rice Diversity
DATA AVAILABILITY
however, several members of evangelical churches still cultivate rice,
including black rice, for home consumption and sale.
In spite of half a century of policy recommendations to change
or abandon their traditional agriculture, Maroon rice farming has
shown a remarkable resilience. Commercial rice entered their diet
decades ago (Geijskes, 1955; Bilby et al., 1989), but did not replace
their own landraces. New waves of Ndyuka Maroons arrived in
French Guiana as refugees during the Surinamese civil war in the
1980s, which led to a rapid urbanization of the region around St.
Laurent du Maroni and the abandonment of agriculture (Fleskens
and Jorritsma, 2010; Léobal, 2016). This provided more farmland
for those that stayed in their forest villages and created a market
for homegrown rice in St. Laurent. Mechanical rice mills, often
donated in development projects (e.g., Rosebel Gold Mines, 2017;
Conservation International, 2018), can lead to malnutrition and
thiamine deficiency as they remove the nutritious bran and germ
of the rice (Lanska, 2010; Ryan, 2011). But Maroon farmers are
well aware of this, and have not given up the traditional dehusking
methods with mortar and pestle.
The Maroons, however, do not have to remain solely responsible
for protecting their cultural heritage. The fascinating history of
Maroon rice, its unique diversity, dynamic character, distinctive
taste and strong gender dimension (which is reflected in varieties
known as ‘dancing woman’, ‘beautiful woman’ and the names
of female ancestors) offer opportunities for broader culinary
marketing and ‘agro-ecotourism’ (see Maxted et al., 2002). The
general public in the Guianas is hardly aware of Maroon rice. Not a
single restaurant in Cayenne or Paramaribo sells Maroon rice dishes.
Given the increased interest by global consumers in traditional food
products (Ardenghi et al., 2018), a greater awareness of the unique
Maroon rice varieties could stimulate their conservation in the face
of increased urbanization and outside pressures. An appreciation of
the role of Maroons as custodians of rice diversity would benefit not
just local food security, but also safeguard a precious global resource.
The datasets generated for this study can be found in Zenodo,
10.5281/zenodo.3276199.
AUTHOR CONTRIBUTIONS
TA, HM, and HB conceived and designed the study; DL and TP
gave input on the study design; TP and TA organized fieldwork;
JA provided the data collection format, germplasm storage, and
phenotyping data; TA and AB carried out the fieldwork; AB,
MV, VM, and HB carried out the genetic research. All authors
reviewed and approved the final manuscript.
FUNDING
This research was funded by the National Geographic Society
(GEFNE grant no. 18416), Naturalis Biodiversity Center (TA), and
the Van Eeden fund (AB). VM was supported by the Marie Curie
Actions of the 7th European Community Framework Programme:
FP7-MCA-ITN 606895 MedPlant. MV was supported by European
Union’s Horizon 2020 research and innovation programme H2020
MSCA-ITN-ETN 765000 Plant.ID.
ACKNOWLEDGMENTS
We are grateful to the staff of SNRI/ADRON in Nickerie, the
Herbier IRD in Cayenne, and to Frédéric Blanchard (Collectivité
Territoriale de Guyane, Cayenne), Brian Mawdo, Manon
Plasschaert, Beatrice Rostand, and Lani Pesna for facilitating our
fieldwork. We thank Frans Afi, Stieven Scheinemann and Edith
Adjako for their work as translators. We are indebted to rice
farmers Maneska Manu, Martha Afonsoewa, Cynthia Asoiti, Elise
Sandiana, Nesia Atanso, Christine Audo, Samantha Dansman,
Christine Feno, Comina Eeswijk, Eleni Galimo, Sonia Sini, Ajadie
Abetau, Ifna Asaida, Marceline and Vanessa Kodeli, Jomea Nyanfai,
Agnes Awenkina, Sylvie and Maloe Deel, Leonie Altret, and Paisie
Mbola for sharing their knowledge and their rice varieties with
us. We thank Sally Price for sharing her unpublished data on
Saramaccan rice varieties. Marlies Lageweg designed the poster
of the different rice varieties. Linh Nguyen Nhat helped us with
the lab work at DNA lab of the Natural History Museum of the
University of Oslo (UiO), Norway. This work was performed on the
Abel Cluster, owned by the University of Oslo and the Norwegian
metacenter for High Performance Computing (NOTUR), and
operated by the Department for Research Computing at USIT, the
University of Oslo IT-department, http://www.hpc.uio.no/.
LINKS TO VIDEO FOOTAGE MADE ON
MAROON RICE CULTIVATION AND
PROCESSING
VIDEO 1 | Two ways of planting rice: either in small holes (‘diki olo’) dug with a
hoe (‘tyap’), or cast over a cleaned field (‘fringi’) after which the rice is covered with
loose soil and pressed flat. https://www.youtube.com/watch?v=rWUDuZCKFr0.
VIDEO 2 | Harvesting rice with a small knife in St. Laurent du Maroni, French
Guiana. This video also shows how interviews were carried out in the field.
https://www.youtube.com/watch?v=bQ1tyfhXIJI&t=110s.
VIDEO 3 | Traditional way of threshing rice by Maroons in French Guiana. https://
www.youtube.com/watch?v=srwL5GrLLW4.
VIDEO 4 | Traditional dehusking of rice by Maroons in Suriname. https://www.
youtube.com/watch?v=KWXIXHhhf4g&feature=youtu.be.
SUPPLEMENTARY MATERIAL
VIDEO 5 | Traditional winnowing of rice with a wooden tray in Suriname. https://
www.youtube.com/watch?v=_5dbhOnqAMU&feature=youtu.be.
The Supplementary Material for this article can be found online
at: https://www.frontiersin.org/articles/10.3389/fpls.2019.01161/
full#supplementary-material
VIDEO 6 | How the Maroon ancestors hid rice grains in their hair. https://www.
youtube.com/watch?v=4H1IbY6PGIk.
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September 2019 | Volume 10 | Article 1161
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Maroon Rice Diversity
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Conflict of Interest Statement: The authors declare that the research was
conducted in the absence of any commercial or financial relationships that could
be construed as a potential conflict of interest.
Copyright © 2019 Van Andel, Veltman, Bertin, Maat, Polime, Hille Ris Lambers,
Tjoe Awie, De Boer and Manzanilla. This is an open-access article distributed
under the terms of the Creative Commons Attribution License (CC BY). The use,
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September 2019 | Volume 10 | Article 1161