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Ethnobotanic and genetic diversity
of sugarcane in French Polynesia
Marotea Vitrac, Taivini Teai, Ines Shili-Touzi, Jean-François
Butaud, François-Régis Goebel, Catherine Hervouet, Angélique
D’Hont
Correspondence
Marotea Vitrac1*, Taivini Teai1, Ines Shili-Touzi², Jean-François Butaud3, François-Régis Goebel4, Catherine Hervouet5,
Angélique D’Hont5
1Research mixt unity about Insular ecosystems in Oceania (UMR 241 EIO), University of French Polynesia (UPF), Punaauia,
French Polynesia
2Agro development engineer school, ISTOM (UR ADI-Suds), Angers, France.
3Independent botanist, Tahiti, French Polynesia & Correspondent of the National Museum of Natural History, Paris, France.
4Agroecology and sustainable Intensification of annual crops, CIRAD, University of Montpellier, Montpellier, France.
5CIRAD, UMR AGAP Institut, Montpellier F-34398, France, UMR AGAP Institut, Université Montpellier, CIRAD, INRAE, Institut
Agro, Montpellier, France.
*Corresponding Author: maroteav@gmail.com
Ethnobotany Research and Applications 29:8 (2024) - http://dx.doi.org/10.32859/era.29.8.1-15
Manuscript received: 12/01/2024 - Revised manuscript received: 02/05/2024 - Published: 03/05/2024
Research
Abstract
Background: In French Polynesia, traditional tō (Saccharum officinarum) have been re-exploited in the recent years to
produce organic certified rum. Former botanists have described the sugarcane which were spread by Polynesians during
their migrations of the Eastern Pacific. One of them, referred by botanists as Otahiti was the main cultivar grown for sugar
production until the 1880s.
Methods: Between 2013 and 2017, we collected 15 sugarcane accessions in the Society Islands and examined their taxonomic
status to establish the correspondence with those described by former botanists. Nine morphological traits were repeatedly
measured including stalk colour, tillering, stalk height, stalk diameter, and internode lengths. We also analyzed them using
flow cytometry, PCR markers and in one case molecular cytogenetics.
Results: The results showed 4 modern hybrids cultivars, 9 traditional S. officinarum and one intergeneric hybrid between S.
officinarum and the wild genus Miscanthus floridulus, Tō 'ā'eho. Among the traditional S. officinarum sugarcane cultivars,
we suggested that Polynesian sugarcane called Tō ‘irimotu and Tō re’are’a could correspond to Otahiti.
Conclusions: The studies of processing characteristics also revealed the high Brix of the Tō 'ā'eho and its potential for
producing rum. These types of hybrids support the hypothesis of Pacific being a satellite center of sugarcane diversity.
Key words: Saccharum officinarum, noble sugarcane, Saccharum maximum, French Polynesia, Otahiti
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Background
In the past ten years, interest in sugarcane production has risen in French Polynesia. Owners and producers are organizing
themselves under quality standards such as certified organic agriculture and a recognized geographic origin. The high price
of sugarcane production locally doesn’t allow for economically viable raw sugar production and the only interest is for high-
value rum production (Vitrac et al. 2022). One of the main specificities is the use of local noble Saccharum officinarum
cultivars which are required for the rum to be certified as a “rum of French Polynesia”.
French Polynesia was populated between year1000-1300 before Christ by Polynesians coming from the Western Pacific
(Wilmshurst et al. 2010). In their canoe expeditions from island to island, Polynesians brought with them several plants such
as coconuts, bananas, yams, taro, breadfruits and sugarcane. Sugarcane are generally called tō (Henry 1928; Whistler 2009)
and were used in medicine, ceremonies (Ellis 1831; von den Steinen 1898), as a carbohydrate resource (Lincoln 2020) and to
sweeten food (Vitrac et al. 2018a). It was a plant of major importance for Polynesians, and they had probably selected and
disseminated several cultivars for specific purposes prior, during and after their migrations from island to island over the
years.
When the first European navigators arrived in the French Polynesian islands, starting in the late eighteenth century following
native Polynesian’s knowledge, botanists gave some descriptions of the S. officinarum cultivars and closely related wild
species (S. maximum and Miscanthus floridulus) they discovered (Wray 1853).
In Tahiti, between 8 to 10 cultivars (Table 1) were described by former botanists such as Cuzent (1860), Nadeaud (1873) and
Henry (1928) and 14 in the Marquesas (Brown 1931) with no correspondence in the Polynesian names due to the different
languages used between archipelagos. One of the S. officinarum cultivars was called “Otahiti” (Bougainville 1771) without
any reference to its Tahitian name and the European explorers spread it all around the world, where it became the main
crop of S. officinarum for sugar production until the years 1880s (Stevenson 1965) because of its high sugar production rate,
robust growth, and excellent ratooning.
At the beginning of the twentieth century, noble S. officinarum cultivars used for industrial purposes were replaced
worldwide initially by intraspecific hybrids and later by interspecific hybrid between S. officinarum and S. spontaneum (herein
“modern cultivars”) developed by breeders, which were more resistant to disease and with a better yield for sugar
production (Stevenson 1965).
As sugarcane was reproduced using setts (stalk cuttings) and without any rules regarding world plant exchanges, many S.
officinarum from different regions and modern cultivars were spread, particularly in all the Pacific islands, following the
navigator’s travels. In French Polynesia, the Otahiti cane was used for sugar production until the end of the nineteenth
century (Ellis 1831; Nollinberger 1857; Cuzent 1860) and the decline of Atimaono plantation in Tahiti (Vitrac et al. 2015). The
Otahiti cultivar, and indeed all noble S. officinarum cultivars, were no more used for sugar or rum production (Fahrasmane
& Ganou-Parfait 1997), after this period.
Rum producers today hope that growing traditional Polynesian S. officinarum cultivars, including cultivar Otahiti, can
enhance the marketability and branding of local products. However, in such a context, it is difficult to be sure which of the
cultivar used today can be regarded as Otahiti sugarcane.
The objectives of this study were to i) collect, record and characterized the diversity of Saccharum and its close relatives
currently present in French Polynesia, with a special focus on S. officinarum; ii) to try and make correspondence between
the accessions we collected and those described by former botanists. To this end, we used morphological descriptions,
process indicators and molecular tools.
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Table 1. Taxonomy and descriptions of the sugarcane cultivars and associated wild species in Tahiti Island by former
botanists
Cuzent (1857, 1860)
Henry (1928)
Scientific name
Tahitian name
main characteristics
Tahitian
name
main characteristics
Saccharum officinarum
Tō ‘irimotu
green color, brittle skin,
urticating hairs
Tō ‘irimotu
purple with brittle skin
Saccharum officinarum
Tō rurutu
Tō rutu
slightly purple and purple
leaves
Tō tea
Tō rutu
light greenish-yellow
Saccharum officinarum
Tō pi’avare /
pi’avere
slightly red, small
internodes, small
diameter
Tō pi’avare
/ pi’avere
grape colored, white
bloom
Saccharum officinarum
Tō ‘ōura / ‘o ‘ura
purplish with yellow
bands, large diameter
Tō ‘ōura /
‘o ‘ura
striped green, white, and
purple
Saccharum officinarum
Tō ‘ute
violet skin and medulla,
big stem, lots of juice,
imported from Batavia by
Bougainville
Tō 'ute
dark red
Saccharum officinarum
Tō Vaihī
Tō Vaihī-‘uo’uo
Tō ‘uo’uo
white, lots of sugar,
imported from the
Sandwich Islands
Tō ha’avai
no description
Saccharum officinarum
Tō ‘avae
yellowish, green stripes
Saccharum officinarum
Tō 'ofe
great light greenish-
drab, resembling a
bamboo
Saccharum officinarum
Tō rā'au
light maroon color, very
hard
Saccharum officinarum
Tō ’ō’opu
very dark purple
Saccharum maximum
Tō ‘ā’eho
green (white), slender
stem, mountain cane,
Tō ‘ā’eho
light green, wild,
resembling a reed
Saccharum maximum
Tō patu
violet (red), slender stem,
mountain cane,
Miscanthus floridulus
'ā'eho
reed
Materials and Methods
Material
Between 2013 and 2017, we prospected the islands of Tahiti, Moorea, Raiatea and Taha’a in the Society archipelago, to
collect all sugarcane and wild species from Saccharum or close genera we could find.
We collected 15 accessions which are described in Vitrac et al. (2018b, 2019a, 2019b) and were tentatively classified as eight
noble cultivars (S. officinarum), four modern cultivars (Saccharum spp.) and two wild accessions (S. maximum and
Miscanthus floridulus) (Table2). In addition, two noble canes, ‘Batavia’ and ‘Black Cheribon’, and one modern cultivar B69566
were imported from the Visacanneâ quarantine (CIRAD, Montpellier), and leaves, from the ‘Lahaina’ noble S. officinarum
from the HARC collection (Hawaiian Agricultural Research Centre). DNA from the modern cultivar R570 was used in
Montpellier for the genetic analysis. The canes were cultivated at farm scale under the same cropping standards described
in Vitrac et al., (2019a) at two separate locations (respectively “collection plot” and “plantation plot” described in Vitrac et
al., (2023)).
DNA fingerprinting and PCR diagnostic markers
DNA extractions were done at the laboratory of the “Epic Vanille” in Raiatea Island and at CIRAD according to the protocol
of Hoisington (1992). DNA fingerprinting with microsatellites was performed at the AGAP institute Genotyping Platform GPTR
of CIRAD (Montpellier, France) as described by Kagy et al. (2016). Six microsatellites markers were used: CV29, CV37 and
CV38 (from Macheroni et al. 2007) and 3 developed by CIRAD (mSScir_14, mSScir_19, mSScir_164). After amplification with
a set of M13 tailed fluorescent labelled primers, fragment sizes were estimated using internal size standard on a DNA
Ethnobotany Research and Applications
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analyser (ABI 3500xL, Applied Biosystems), and allele scorings were analyzed using Genemapper v4.1 software (Applied
Biosystems).
A Miscanthus PCR diagnostic marker was developed from a Miscanthus inter Alu type transposable element sequence (Alix
et al. 1999). Two primers (5’-GTGACTCCTGCTGTGACTCC-3’ and 5’-GAACATGATCGGAGGCCCTC-3’) were designed in the
Miscanthus sequence MsCIR2 (EMBL Y17576, Alix et al. 1999). These primers used on Miscanthus floridulus or Miscanthus
sinense DNA produced of band of around 300 bp and no amplification was observed with Saccharum DNA. The amplification
reaction was performed in a final volume of 25µl, containing 25ng of sugarcane DNA, 0.2mM dNTP mix, 2mM MgCl2, 80mM
Tris-HCl buffer, each primer at 0.1µM and 0.06U of Taq polymerase (FIREPol®). The PCR was carried out with the following
program: 94°C for 4 min; 35 cycles of 94°C for 30 s, 60°C for 45s, 72°C for 1min; a final cycle of 72°C for 10min. PCR products
were separated on 1% agarose gel and visualised after staining with ethidium bromide.
Morphological analysis
Table 2: Sugarcane and wild accessions collected in French Polynesia or imported, and methods used to characterize them.
Year
Names
ref.
Process
Morphology
PCR markers
Flow cytometry
Origin
Saccharum officinarum
collected
2013
Verte à Bandes Pourpres
VBP
x
x
x
x
17°32'01.1"S 149°25'52.1"W
2013
Rouge à Bandes Vertes
RBV
x
x
x
x
17°45'2.80''S 149°21'11.9''W
2013
VErte
VE
x
x
x
x
17°30'47.4''S 149°30'3.18''W
2013
Trois(3) Couleurs
3C
x
x
x
x
17°40'34.4''S 149°18'28.3''W
2013
Jaune à Rayure Pourpre
JRP
x
x
x
x
17°45'37.0"S 149°22'02.7"W
2013
POurpre
PO
x
x
x
x
17°30'53,9''S 149°28'10.3''W
2014
Jaune à Taches Rouges
JTR
x
-
x
-
17°36'59.3''S 149°18'8.04''W
2014
ROuge
RO
x
-
x
x
17°43'44.0"S 149°18'51.1"W
2017
Rapa JFB
-
-
x
-
Rapa island
imported
2017
BATtavia
BAT
x
-
x
x
CIRAD Visacane
2017
Black Cheribon
BC
-
-
x
x
CIRAD Visacane
2017
Lahaina
-
-
x
-
HARC Hawaii
Saccharum spp.
collected
2013
Rouge Reflets Verts
RRV
x
x
x
x
16°44'09.5"S 151°26'19.9"W
2013
Jaune Roseau
JR
x
x
x
x
17°45'56.0"S 149°28'00.3"W
2014
Blanche
Bla
x
-
x
x
17°45'56.0"S 149°28'00.3"W
2014
HAWaii
HAW
-
-
x
-
17°44'60.0"S 149°21'41.0"W
imported
2016
B69566
Ble
x
x
x
x
CIRAD Visacane
2018
R570
R570
-
-
x
-
CIRAD AGAP
Saccharum maximum
collected
2016
Tō 'ā'eho
x
-
x
x
17°49'12.3"S 149°08'06.3"W
Miscanthus floridulus
2016
‘ā 'eho
x
-
x
x
17°37'56.4"S 149°36'48.0"W
x: done ; -: not done
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To compare our plant material to the descriptions made by the former botanists for all the sugarcane observed, we used the
UPOV (2005) sugarcane varietal repository (using Artschwager & Brandes (1958) standards). These sheets are not presented
in this study, being too large (54 traits), but it was the first step of identification and comparison to descriptions given by
former botanists. The main traits described were: stalk color, stalk diameter, stalk height, tillering (number of stalks per
plant), ratoon ability, internodes length, leaf color, leaf bunch appearance and flowering occurrence. We also compare our
observations to the recent descriptions made by Lincoln (2020) who described all the Hawaiian sugarcane.
For nine varieties as described in Table 2 (“morphology” column), ten stalks of each variety and each plot were sampled from
the plantation plots in August 2015 (first plantation, 12 months) and August 2016 (first ratoon, 24 months). The stalk height,
the stalk diameter and the internode length as well as the tillering were measured.
Data was analyzed using the statistical software XLSTAT 19.4.45191. A population probability law (normal distribution) and
descriptive statistical parameters such as means and standard deviations were processed. Mean comparison tests of Mann
Whitney (samples<30) were used to compare stalk height, tillering, stalk diameter, and internode length. A factorial analysis
was also done.
Flow cytometry
Leaves were sampled from both collection and plantation plots and sent fresh to AGAP laboratory to be analyzed by flow
cytometry method accorded to Ochatt (2011) to determine their nuclear DNA content and evaluate chromosome numbers.
Leaves from accessions of Miscanthus sinense were used as standard.
Genomic In Situ Hybridization (GISH)
Genomic hybridization was performed according to D’Hont et al. (1996) on chromosomes of the wild Saccharum maximum
accession Tō 'ā'eho using as probes 200ng of Miscanthus sinense NG7722 genomic DNA (labeled with Biotin and detected
with avidin-Texas Red) and 200ng of S. officinarum BN3066 genomic DNA (labeled with digoxigenin and detected with anti-
digoxygenin-FITC). The chromosomes were counterstained with 4’-6 diamidino-2-phenylindole (DAPI; Vectashield Mounting
Media with DAPI), and fluorescent images were then captured separately using a cooled high-resolution CCD camera (ORCA
Hamamatsu) attached to a Leica DMRXA2 fluorescent microscope.
Process indicators
Three stalks for each cultivar (Table 2) of the collection plots were milled (laboratory stainless steel miller) to determine the
juice yield (weight of juice / weight of milled cane), the fibre content (weight of fibre residue after milling / weight of cane
before milling) and the Brix degree (soluble dry extract, with the ATAGO® “pocket” digital refractometer) according to
traditional milling methods (Rein 2017).
Results and discussions
DNA fingerprinting
DNA fingerprinting of the 15 collected accessions was performed using 6 Saccharum microsatellites markers to determine if
some of the collected accessions could be identical or correspond to somaclonal variants (Figure 1).
For all accessions except the Miscanthus accession, an amplification was obtained. The absence of amplification for the
Miscanthus accession was expected since these markers are specific to the Saccharum genus. Very similar fingerprints were
obtained for ‘Verte bandes pourpres (VBP)’, ‘Verte (VE)’ and ‘Jaune à taches rouges (JTR)’. However, ‘Verte bandes pourpres’
is striped green and purple, while the two others are solid colored. This difference may thus be due to somaclonal variation.
‘Rapa JFB’ and ‘Trois couleurs (3C)’ had also very similar fingerprints as well as ‘Jaune roseau (JR)’ and ‘Hawaii (HAW)’. ‘Rapa
JFB’, ‘Jaune à taches rouges’ and ‘Hawaii’ were not included in further analysis.
Morphological and phenological comparison of cultivars
Eight of the collected accessions and one imported modern cultivar were cultivated at farm scale under the same cropping
standards, firstly to compare them to the descriptions made by the former botanists and secondly to characterize them using
standard/classical descriptors. After two years of cultivation for most of the cultivars in two separate sites, we encountered
some difficulties establishing a “standard” morphologic identification. Environmental factors such as sunlight exposure,
rainfall, and soil fertility have been reported to influence morphological characteristics (Lincoln 2020). Also, the maturation
stage and age of the plant have a large influence. However, some characteristics are both distinctive and relatively consistent,
Ethnobotany Research and Applications
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and it is the combination of all which finally allows us to identify a cultivar with accuracy. This is similar to the approach by
Lincoln et al. (2022), in which 95 morphological traits were recorded but they found that the best separation of cultivar
classifications occurred when using only 19 parameters. Producing the variety sheets (following Artschwager & Brandes 1958
In: UPOV 2005), we observed some discriminant characteristics as it is resumed in Table 4: stalk color, leaf bunch size and
color, stalk diameter, tillering, internode length, flowering, special characteristics, and potential ratoon. We paid particular
attention for morphological characteristics linked to agronomy.
Figure 1. DNA fingerprinting of the 15 collected accessions (performed using 6 Saccharum microsatellites markers).
Saccharum officinarum (green color, collected and dark dark green color, imported) and Saccharum spp. (pink color, collected
and purple color, imported).
We were surprised that most of former botanists did not mention flowering of sugarcane. Moreover Lincoln (2020)
mentioned that he could not consistently observe the canes throughout the year so could not be confident in his
observations of flowering. In our case, we observed that flowering did not occur every year for most of the cultivars. This
criterion helped us to distinguish two morphologically very similar varieties: Tō ‘irimotu (VE, Table 4) which flowers every
year whereas ‘Jaune à rayures pourpres’ (JRP) a cane we propose to call Tō re’are’a (which means Yellow in Polynesian
language) has never flowered since 2014. We did not find mention of this particular cane by Polynesians nor former botanists,
which is very easy to confuse with Tō ‘irimotu because of their similar nodes zone and stalk colour. We found that flowering
and a few additional agronomic characteristics (tillering, size of internode length and ratoon ability) allowed us to make the
distinction.
Table 3 sums up the measurements done in 2015 and 2016 regarding agronomic characteristics for the sugarcane in the
plantation plots. A factorial analysis based on these morphological characteristics is shown in Figure 2. The first axis of the
factorial analysis (explaining 49.31% of variation) separated two groups. Group 1 was composed by ‘Rouge à Bandes Vertes’
(RBV), JRP, VBP, VE, 3C and ‘Pourpre’ (PO), which we previously classified based on morphological characteristics as S.
officinarum and Group 2 was composed by ‘Rouge à Reflets Verts’ (RRV) and JR, which we previously classified as modern
cultivars, and B69566 (a modern cultivar imported from CIRAD visacane). On the second axis (explaining 27.99% of the
variation), JRP (Tō re’are’a) and RBV (Tō ‘ute, Table 4) cultivars are separated from the other noble sugarcane. According to
our literature interpretations, Tō ‘ute variety (which corresponds to ‘Cavengerie’, (Lincoln 2020)) is probably not from an
ancient Polynesian introduction and there is still doubt regarding Tō re’are’a (JRP).
0
0.2
BAT
BC
Bla
B69566
HAW
JR
JRP
JTR
Lahaina
PO
R570
RBV
RO
RRV
3C
Rapa JFB
VBP
VE
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In accordance with literature regarding the morphological differences between noble sugarcane and modern cultivars, Group
1 has poor tillering (2.80 to 8.83 stems par plant), large stalk diameter (2.73 to 3.61cm), small internodes (5.10 to 7.82cm)
and small stalk height (1.02 to 1.58m except JRP 1.89m) and is significantly different (p=0.05) from the Group 2: rich tillering
(5.37 to 15.00), medium stalk diameter (2.61 to 2.99cm), medium internodes (8.56 to 12.37cm ) and big stalk height (1.82 to
1.98). Only the noble sugarcane JRP height is similar to Group 2 heights. Vitrac et al. (2019b) showed that main components
of Polynesian Saccharum officinarum yield are height and tillering. Tō re’are’a (JRP) cultivar seems to be different from all
others from the Group 1 regarding height. It can be an ancient Polynesian introduced variety or introduced from elsewhere
from an unknown area. Without significant differences, Table 3 shows that sugarcane of Group 2 globally increased all their
characteristics from first plantation (2015) to first ratoon (2016) while sugarcane from Group 1 showed the opposite. For our
S. officinarum cultivars, the best agronomic results were obtained in first plantation which is confirmed by Vitrac et al.
(2019b).
Table 3. Results of measured morphologic characteristics for the different studied cultivars.
Cultivars
Stalk height (m)
Tillering
Stalk diameter (cm)
Internodes length (cm)
Group 1
2015
2016
2015
2016
2015
2016
2015
2016
RBV
1.58 ns
1.49 ns
5.27 a
6.80 ac
3.44 c
3.42 ns
7.10 a
6.75 a
JRP
1.88 b
1.89 c
3.03 ns
2.80 ns
3.58 c
3.61 ns
5.89 ab
7.82 ns
VBP*
-
1.20 a
-
8.83 d
-
2.97 c
-
6.19 ac
VE
1.36 a
1.21 a
5.10 a
5.00 a
2.73 a
2.75 a
6.46 a
6.44 a
3C
1.34 a
1.02 ab
6.73 a
5.03 a
2.83 ab
2.79 ab
5.73 b
5.14 b
PO
1.13 ns
1.04 b
4.07 ns
4.93 a
3.53 c
3.22 ns
5.10 b
5.64 bc
Group 2
RRV
1.94 b
1.90 c
7.07 a
15.00 b
2.95 d
2.99 c
8.56 ns
8.97 d
JR
1.88 b
1.98 c
5.37 a
7.37 cd
2.93 bd
2.61 b
12.37 ns
9.06 d
B69566**
-
1.82 c
-
13.33 b
-
2.76 ab
-
10.53 ns
Std.
deviation
0.19 < σ 2015 < 0.34
1.69 < σ 2015 < 4.42
0.23 < σ 2015 < 0.39
1.05 < σ 2015 < 2.52
0.17 < σ 2016 < 0.35
1.47 < σ 2016 < 5.02
0.22 < σ 2016 < 0.79
0.62 < σ 2016 < 1.87
a, b, c, d, ns: Mann Whitney (p< 0.05) results ; * : not cultivated in 2015 ; ** : introduced in 2016
Figure 2. Factorial analysis following morphological characteristics of the varieties cultivated in the plantation plot.
Saccharum officinarum (green color) and Saccharum spp. (pink color, collected and purple color, imported).
VE
3C
PO
JRP
RBV
VBP
B69566
RRV
JR
-2
-1,5
-1
-0,5
0
0,5
1
1,5
2
2,5
3
-2 -1,5 -1 -0,5 00,5 11,5 22,5 3
Factorial analysis: (axis 1 and 2, 77.31 %)
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Chromosome number estimation through flow cytometry measurement
To further confirme our classification of the collected accessions in S. officinarum versus modern hybrid cultivars, we
analyzed them with flow cytometry. Flow cytometry allows to determine nuclear DNA content which can be related to
chromosomes numbers. S. officinarum typically has 80 chromosomes with some small variation and a nuclear DNA content
of around 7.6 picograms while modern cultivars have around 110-120 chromosomes with a nuclear DNA content of around
10 picograms (D’Hont and Glaszmann 2001). The results presented in Figure 3 and Table 4 showed that the 7 collected
accessions that we suspected based on morphological characteristic to belong to S. officinarum, as well as the two standard
we used for S. officinarum (‘Batavia (BAT)’ and ‘Black Cheribon (BC)’), all have nuclear DNA content typical of S. officinarum.
The three collected accessions that we suspected based on morphological characteristic to be modern cultivars, as well as
the two standards we used for modern cultivars (R570 and ‘Bleue (Ble) B69566’), all have nuclear DNA content typical of
modern cultivars.
Figure 3 Flow cytometry results of S. officinarum versus modern hybrid cultivars and wild species.
Characterization of the Saccharum maximum accession
Former botanists have noted the presence of atypical sugarcane in mountains that they related to Saccharum spontaneum
(Cuzent 1860; Guillemin 1837). The origin of these types of accession have long been debated, having been related to S.
spontaneum, Erianthus or Miscanthus, and they were attributed different taxonomic names, the most commonly used
currently being Saccharum maximum (Price and Daniels 1968; Grassl 1946). We found one such accession, call Tō ‘ā’eho in
Polynesian language. In Polynesian language, ‘ā’eho is the name for Miscanthus floridulus and tō the name for S. officinarum.
Tō ‘ā’eho thus signifies a possible hybrid between ‘ā’eho and Tō. This together with absence of S. spontaneum in Society
Islands suggested that Tō ‘ā’eho could be an intergeneric hybrid between S. officinarum and Miscanthus floridulus.
To further verify this hypothesis, we used microsatellite markers specific to the Saccharum genus and a PCR markers specific
to Miscanthus genus. Both types of markers produced an amplification strongly suggesting that this accession is a hybrid
between the two genera.
We also performed a hybridization on the chromosomes of Tō ‘ā’eho with one probe consisting of total DNA of S. officinarum
(detected in green) and one probe of total DNA of Miscanthus floridulus (detected in red). The results presented figure 4
clearly show that Tō ‘ā’eho is an intergeneric hybrid with around 40 chromosomes inherited from S. officinarum and 19
0
2
4
6
8
10
12
R570
Batavia
Black cheribon
VE
RO
3C
VBP
RBV
PO
JRP
RRV
Bla
JR
Ble
Tō ‘ā’eho
Misc
Nuclear DNA content (pg)
Cultivar
S. officinarum
S. spp
S. maximum
Miscanthus
standards
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9
inherited from Miscanthus. Price & Daniels (1968) analyzed an accession, Raiatea 1, collected in Raiatea Island in 1935 and
based on the observation 2n = 60 chromosomes, they suggested that it could derived from hybridization between Saccharum
and Miscanthus. These results also confirmed, Grassl (1946) hypothesis of the involvement of Miscanthus in Saccharum
maximum.
Figure 4. Chromosome preparation of Tō ‘ā’eho accession after in situ hybridization
with Miscanthus floridulus total DNA (detected in red) and S. officinarum total DNA
(detected in green).
Processing characteristics of the studied accessions
Figure 5 resumes the measurements done (2015 and 2016 as a mean) regarding processing characteristics about cultivated
cultivars in the collection plot.
Figure 5. Processing characteristics (Brix degree, fiber rate and juice yield) of the studied accessions. Purple: introduced
modern hybrid cultivars; pink: collected modern hybrid cultivars; red: Noble Saccharum officinarum introduced from
visacane®; Green: collected noble Polynesian Saccharum officinarum; Blue: wild accessions
The fibre content was almost the same for both S. officinarum and modern hybrid cultivars with around 30-35% (Figure 5).
For Saccharum maximum Tō ‘ā’eho, this rate is about 50% and for Miscanthus floridulus 60%. Also, the juice rate is about
60-65% regarding both S. officinarum and modern hybrid cultivars even for the introduced cultivars (B69566 and Batavia).
So, these process indicators are not relevant here to distinguish S. officinarum from modern hybrid cultivars with laboratory
material we used. Vitrac et al. (2019b) found the opposite with more effective juice extraction for noble S. officinarum
sugarcane. They also found no difference regarding Brix degree, which is richer in modern hybrid cultivars in our study,
although classically Brix is higher in modern cultivars. These discrepancies may be coming from the milling equipment and
refractometer we used and thus not relevant here for taxonomic distinction between S. officinarum and modern hybrid
cultivars.
However, the wild species Saccharum maximum (Tō ‘ā’eho) and Miscanthus floridulus displayed large differences: Tō ‘ā’eho
has 40% juice extraction yield which is 20% less than sugarcane and a valuable Brix degree of about 15, comparable to most
sugarcane cultivars. Tō ‘ā’eho can thus be regarded as an interesting variety for rum production because of its high sugar
content.
5
7
9
11
13
15
17
19
21
10%
20%
30%
40%
50%
60%
70%
80%
Bla
B69566
JR
RRV
BAT
VE
JTR
VBP
PO
3C
RO
RBV
JRP
To'a'eho
Misc
Brix degree
Fiber rate (%) Juice Yield (%) Brix degree
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Table 4: Identification and characterization of the sugarcane and wild species of the study.
Species
origin
Polynesian
name
ref.
common
name
stalk color
leaf
bunch
leaves
diameter
tillering
internodes
flowering
specials
ratoon
nuclear
DNA
content
(pg)
S. officinarum
F.P.*
Tō ‘irimotu
VE
verte
green
falling
back
large,
green
large
3 to 10
small
every time
brittle
skin
1 to 3
7.66
S. officinarum
F.P.
Tō rutu
RO
rouge
red
falling
back
large,
purple
large
3 to 6
small
every time
-
0
7.56
S. officinarum
F.P.
Tō pi’avare
3C
trois couleurs
red, brown and
yellow
falling
back
large,
green
small
3 to 10
small
sometime
-
1 to 3
7.66
S. officinarum
F.P.
Tō ‘ōura
VBP
verte à bandes
pourpres
green with
abundant large
purple stripe
falling
back
large,
green
large
3 to 10
small
no
-
1 to 3
7.56
S. officinarum
F.P.
Tō ‘ute
RBV
rouge à bandes
vertes,
Cavengerie
red with large low
abundant large
purple stripe
falling
back
large,
green
large
3 to 10
large
rare
-
1 to 5
7.65
S. officinarum
F.P.
Tō ’ō’opu
PO
pourpre, Badila
very dark purple
falling
back
large,
green
large
3 to 10
very small
sometime
-
1 to 3
7.58
S. officinarum
F.P.
Tō re’are’a**
JRP
jaune à rayure
pourpre
yellow with
abundant very
thin purple stripe
falling
back
large,
green
large
3 to 6
small
no
-
1 to 5
7.59
S. officinarum
CIRAD
-
BAT
Batavia
yellow with
abundant large
purple stripe
falling
back
large,
green
large
3 to 10
small
no
-
1 to 3
7.52
S. officinarum
CIRAD
-
BC
Black Cheribon
purple
falling
back
large,
green
large
3 to 10
small
no
-
1 to 5
7.51
S. maximum
F.P.
Tō ‘ā’eho
Tō
‘ā’eho
-
green, brown and
red
erected
thin,
green
very small
10 to 30
large
every time
pubescent
> 5
6.46
M. floridulus
F.P.
‘ā’eho
‘ā’eho
-
green
-
thin,
green
very small
10 to 30
large
every time
-
-
4.62
Saccharum spp.
unknown
-
RRV
Rougereflets
verts
red as purple
falling
back
large,
green
large
7 to 20
large
no
green
stripe
(rare)
> 5
10.05
Saccharum spp.
CIRAD
-
Bla
blanche
white
erected
thin,
green
small
7 to 20
large
everytime
cerusia
> 5
9.96
Saccharum spp.
Unknown
-
JR
jaune roseau
yellow
erected
thin,
green
small
7 to 20
large
no
cerusia
> 5
11.07
Saccharum spp.
CIRAD
-
Ble
B69566
blue
falling
back
thin,
green
small
7 to 20
large
no
cerusia
> 5
-
Saccharum spp.
CIRAD
***
R570
-
-
-
-
-
-
-
-
-
-
10.69
* : French Polynesia ; ** : no descriptions in literature ; ***: not cultivated, from literature
Ethnobotany Research and Applications
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Discussion
In this study we characterized the Saccharum diversity we could collect in part of the Society Islands (Vitrac et al. 2019b),
tried to make correspondence between theses accessions and the one described by former botanists, and tried to distinguish
traditional S. officinarum Polynesian cultivars from more recently introduced S. officinarum and modern hybrid cultivars.
Sugarcane has been propagated vegetatively through cutting by farmers since their arrival on these Polynesian islands some
1000 years ago. Over this period, mutations occurred through somaclonal variations which when favorable or attractive
(bright color for example) could have been selected by farmers. In addition, although S. officinarum is generally not very
fertile, it can sometimes flower when environmental conditions are favourable and result in some progenies from which
farmers could also select attractive clones. The occasional generation of somaclonal variants, the frequent exchange of
material between islands often associated with a change of Polynesian names and the frequent mix-up of materials make
correspondence between accessions over time or regions quite difficult. It is Even more difficultto differentiate accessions
introduced originally by the first Polynesian from more recently introduced ones. Lincoln et al. (2022) have reported on these
difficulties while analyzing the sugarcane diversity in Hawaii and highlight the need to combine several lines of evidence
including, morphological characteristic, genetics characteristics as well as ethnological and historical research.
We collected 7 Saccharum officinarum, 1 Saccharum maximum, 3 modern hybrid cultivars Saccharum spp. and 1 Miscanthus
floridulus (Table 4). These numbers are very close to what former botanists described (Cuzent 1860; Nadeaud 1873; Pancher
1855; and Henry1928). Among the 7 S. officinarum, a few had identical fingerprints, some of them representing somaclonal
variants with clear distinct morphological characteristics.
Regarding the modern cultivar Saccharum spp. group which were introduced in French Polynesia before 2013, we have few
information about them, and it was impossible to know their true origin. The ‘Blanche’ (Bla) variety came from CIRAD and
was introduced in the 1970s. At the same period, the JR variety was introduced maybe from Australia or from Hawaii as well
(personal information from a sugarcane producer). We don’t have any information about the RRV.
Some S. officinarum described in old literature (Table 1) as Tō ‘avae, Tō rā’au, Tō vaihī, Tō ha’avai, Tō ‘ofe were not found,
observed or identified during our study and fieldwork. They may have disappeared because not cultivated anymore or
difficult to link to the ones we collected because of traits variation due to environment or maturation stage when observed.
We can also note some contradictions and oddities between sources: for example, according to Cuzent (1860), Tō rutu
(‘Rouge’ (RO)) has red stalk contrary to Henry (1928) observation as green. Also, for Henry (1928) Tō ‘irimotu (VE) is purple
contrary to Cuzent (1860) observation as green. But maybe they were talking of the same variety for which color may depend
on conditions. For example, the one we called Tō re’are’a (JRP) can be green or yellow and sometimes with abundant thin
purple stripes.
Lincoln (2020) mentioned around 50 traditional Hawaiian Saccharum officinarum, 12 introduced S. officinarum and 7
recognized modern cultivars. This result was supported by Schenk et al. (2002) genetic diversity study who found 41 old/
traditional S. officinarum with several of them being clustered together in genetic analyzing suggesting that they differed by
somaclonal variations. Moreover, Wilfong in 1883 found 50 varieties of ancient Hawaiian introduction. It is surprising to see
that former botanists mentioned no more than 8 to 14 S. officinarum varieties in French Polynesian islands as opposed to
around 50 in Hawaii. We would have expected the opposite since Polynesians first arrived in Marquesas and Society islands
and then went to Hawaii. The diversity of native varieties in Hawaii that arose most probably through somaclonal variation
is likely driven by the importance of sugarcane in the pre-European context in Hawaii, as well as the ecological diversity that
varieties would have needed to be selected for. In addition, we can suppose that many more introductions of noble varieties
from other pacific islands or countries occurred in the 18th century in Hawaii, before the creation of the breeding stations.
It is to be noted that Saccharum maximum was not found in Hawaii (Wagner et al. 1999) and is known in Marquesas, Society,
Austral and Cook archipelagos.
Although, we can find some contradictions in historical records, they can help distinguish accessions introduced originally by
the first Polynesian from more recently introduced ones.
The sugarcane multiplication is vegetative so, following Polynesian migrations from Tahiti and Marquesas Islands to Hawaii,
we would expect to find there most of Marquesan and Tahitian varieties in Hawaii. It seems to be the case based on
morphological characteristics for Tō ‘ute (‘ie’ie, ‘Cavengerie’ or RBV), Tō ’ō’opu (‘Badila’ or PO) according to Lincoln (2020)
but not regarding other noble varieties. However, we can note some similitudes between Tō rutu (RO) and ‘Honua’ula’ which
Ethnobotany Research and Applications
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is the only Hawaiian sugarcane that has purple leaves too. Tō ‘ōura (VBP), green with big purple stripes, is similar to ‘Not
Laukona’ and ‘Hawaiian officinarum’, and of uncertain origin and evocated by Wray (1853) as an introduced sugarcane in
Tahiti by Bougainville from Batavia (current Jakarta) in 1769, the molecular fingerprint showing the opposite (VBP very close
to VE).
Regarding Tō ‘ute (RBV) and Tō ’ō’opu (PO). These canes are described by Lincoln (2020) as introduced Hawaiian S.
officinarum. Tō ‘ute is called Kō ‘ie’ie or Cavengerie (Kō is Tō in Hawaiian language) and was introduced in Hawaii from New
Caledonia while it was introduced in Tahiti from Batavia Island (Cuzent 1860). However, we can note some contradictions as
Lincoln (2020) describe ’ie’ie with a small diameter and poor ratooning which is contrary to Tō ‘ute from Tahiti.
Tō ’ō’opu (PO), imported in the 1920s in Hawaii, is called Badila and is one of the important S. officinarum used in breeding
programs to produce Saccharum spp. modern cultivars, like Black Cheribon and ‘Lahaina’ (Heinz 1987).
Lahaina is an another introduced variety in Hawaii, which is recognized as Otahiti cane and coming from Marquesas Islands
in 1853 (Wilfong 1883). It is surprising to see that Brown (1931) doesn’t mention any Otahiti cane in the Marquesas islands.
He actually mentions 14 sugarcane in the Marquesas with no name correspondence with names collected in the Society
islands.
For Nadeaud (1873), Tō ‘uo’uo, vaihī and rurutu (rutu, RO) are imported and for Cuzent (1860) only Tō ‘ute (RBV) was
imported following Wray (1853). In a letter from Pancher (former botanist of Tahiti and New Caledonia) to his colleague
Decaisne at the beginning of 1855 (unpublished data), some more important indications can help us regarding identification
of noble Polynesian canes: the importation from Sandwich Islands (current Hawaii Islands) of avahi variety is mentioned and
seems to be accurate. Pancher indicates that this variety avahi (different spelling of vaihī or ha’avai) is named ’uo’uo by the
young Tahitians and we then conclude that Tō ha’avai (Henry 1928), vaihī, ‘uo’uo (Cuzent 1860) and avahi are the same
variety which doesn’t exist anymore nowadays. He also mentions that Tō pi’avare (3C) and Tō ‘irimotu (VE) are among the
oldest Polynesian introduced varieties. Tō pi’avare (3C) which is recognized as a medicinal cane (Vitrac et al. 2018a) is often
confounded with other sugarcane. For example, in a recipe for traditional medicine, Hooper (1995) mention Tō patu from
Raiatea (described as a wild species in Tahiti by Cuzent, 1860 and Nadeaud, 1873) as Tō pi’avare from Tahiti, vernacular
names being different in a single archipelago. It is also maybe called Tō rā’au by Henry (1929) because of its medicinal use
(the word rā’au in Polynesian language meaning a medicine), the description corresponding to the observations we made
but Tō rā’au being actually Tō pi’avare (3C). Lincoln (2020) documents how in Hawaii, names for certain cane varieties were
tied to their usage, and it is possible that certain names were applied to multiple different cane varieties when they were
used in a particular way.
Cuzent (1860) says that Otahiti sugarcane cultivated in the Caribbean French territories has a green or a yellow stripe, which
is only the case regarding Tō ‘irimotu (VE) and Tō re’are’a (JRP). Cuzent (1860) and Henry (1928) also mention that cultivated
fields presented flowers from May to July without any other distinctions. We then can suppose that the main cultivated
sugarcane were the Otahiti ones, with green or yellow stripe, which is only the case for Tō ‘irimotu (VE) and Tō re’are’a (JRP)
and as only Tō ‘irimotu (VE) is flowering every year (and Tō re’are’a (JRP) not), we can suppose that Otahiti could be Tō
‘irimotu (VE).
Tō re’are’a (JRP) is one of the most productive noble sugarcane cultivated in Tahiti (Vitrac et al. 2019b) and looks very similar
to Yellow Caledonia described by Lincoln (2020) which is an introduced famous productive cane from New Caledonia.
However, Lincoln (personal discussions) says that Yellow Caledonia cultivar is definitely different from Tō re’are’a (JRP).
We finally can ask the question of Otahiti cultivar as a single variety or group of varieties? Wilfong (1883) and Lincoln (2020)
say that Lahaina is the Otahiti sugarcane introduced in Hawaii in 1853 from the Marqueseas. Tō ’irimotu (VE) seems to be
this Otahiti sugarcane, but we also can note that Tō re’are’a (JRP) is morphologically very similar to Tō ’irimotu (VE), the
difference being the flowering. We then can suppose that Tō re’are’a (JRP), if from ancient Polynesian introduction, is also a
probable Otahiti sugarcane, especially because of its higher agronomic and sugar productivity than other Polynesian S.
officinarum cultivars.
In consideration of all these points, the Polynesians sugarcane regarded as having been introduced by ancient Polynesians
are: Tō ’irimotu (VE), Tō pi’avare (3C), Tō rutu (RO) and Tō ‘ōura (VBP). Tō re’are’a (JRP) is not described in old literature or
has been confounded with another cultivar. The 19th century introduced sugarcane are: Tō ‘ute (RBV) and Tō ’ō’opu (PO).
All the modern Saccharum spp. cultivars are recently introduced (>1900, date of the very first hybridization stations): (i) with
unknown origin: rouge reflets verts (RRV), jaune roseau (JR), and (ii) with known origin: Blanche (Bla).
Ethnobotany Research and Applications
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Conclusion
Tahitian sugarcane cultivars were not kept in a collection and have not been in cultivation for long, this lead to difficulties in
recording of their names and origins i.e. introduction or not.
In this study, we were able, using agro-morphological traits and molecular tools, to distinguish, among the 15 collected
accessions, we recently collected, the ones corresponding to S. officinarum, modern hybrid cultivars and Saccharum
maximum. We could characterize 7 distinct S. officinarum accessions/cultivars.
We were also able to tentatively make correspondence between the collected S. officinarum and the one reported in ancient
literature. The morphological approach suggested that original Otahiti could be Tō ‘irimotu (VE) or Tō re’are’a (JRP). Some
distinctive points such as flowering (no flowering for Tō re’are’a) indicate that Tō ‘irimotu could be the probable Otahiti
described and exported around the world by Bougainville. On the other hand, Tō re’are’a could also be the probable Otahiti
because of its high agronomic potential that must have attracted former farmers.
We also confirmed that the wild species Saccharum maximum is a hybrid between S. officinarum and Miscanthus floridulus
and revealed that it has good Brix degree and juice production to produce rum.
In French Polynesia, the characterization of the current cultivars will help significantly the sugarcane and rum producers to
be sure of what they are growing and to keep safe their recognized geographic origin label that imply the use of the
traditional Polynesian S. officinarum cultivars.
Declarations
List of abbreviations: AGAP: Amélioration génétique et adaptation des plantes méditerranéennes et tropicales; BAT: Batavia;
BC: Black Cheribon; Bla: Blanche ; Ble: Bleue; CCD: Cooled high-resolution Camera; CIRAD: Centre International pour la
Recherche Agronomique et le Développement; DAPI: Vectashield Mounting Media; DNA: Desoxyribo Nucleic Acid; GISH:
Genomic In Situ Hibridization; HARC: Hawaiian Agricultural Research Center; JFB: Jean François Butaud; JR: Jaune Roseau;
JRP: Jaune à Rayure Pourpre; JTR: Jaune à Taches Rouges; PO: Pourpre; PCR: Polymerase chain reaction; RBV: Rouge à Bandes
Vertes; RO: Rouge; RRV: Rouge Reflets Verts; S.: Saccharum; Spp.: species; Std.: Standard; UPOV: Union internationale pour
la protection des obtentions végétales; VBP : Verte à Bandes Pourpres; VE: Verte HAW: Hawaii; 3C: Trois Couleurs ;
Ethics Approval: To respect the ISE Code of Ethics Guidelines, before any field sampling, we first got the authorization of
local authorities (“Direction de l’Agriculture” of French Polynesia) and secondly ask the agreement of every people where
we found the sugarcanes before collect.
We everytime to everybody explained our objectives as first renew knowledge about ancient sugarcanes used by local people
and secondly help to develop organic rum production by a recognized geographic origin procedure.
We kept cuttings of all varieties and create some collections in the local administrations (“Direction de l’Agriculture”) and
museums (“Musée de Tahiti” and “Fare Natura”). We also expect to export them in a quarantine center to protect them
(CIRAD visacaneÒ).
We also will produce a poster showing the list of cultivars found with their Polynesian names, exposed in these special places.
We finally will publish the list and the characteristics of the cultivars in the local scientist journal which name is “bulletin de
la société des océanistes”.
Consent for publication: All authors consent for publication
Availability of data and material: All data and material can be produced in transparency.
Competing interests: The Authors declare no competing interests.
Funding: The research was supported by both CIRAD and University of French Polynesia.
Author’s contributions: Mr Vitrac Marotea realized the experimentations, he drove the study and wrote the paper; Mr
TeaiTaivini contributed to the aim of the study and the global design; Mrs Shili-Touzi Ines contributed to the writing of the
paper and corrections as well; Mrs Hervouet Catherine done the genetic analysis and contribute to the writing of the paper;
Mr Butaud Jean-François contribute to the morphologic observations and interpretations. He also corrected the paper; Mr
Goebel Francçois-Régis drove the paper and contribute to original design with statistics; Mrs D’Hont Angélique drove the
paper as well and contribute to the global design. She drove the genetic items and contribute to write the paper.
Acknowledgments
We thank Susan Schenk, former researcher at the HARC who send us the leaves of Lahaina in 2016 and Noa Kekeuewa Lincoln
(Botanist researcher, UH Manoa, Hawaii) for his help identifying the varieties common between Hawaii and Tahiti.
Ethnobotany Research and Applications
14
We also thank Sandra Lepers Andrzejewski (genetic researcher) for the DNA extractions done at the Epic Vanille de Tahiti of
Raiatea laboratory.
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