Planta (2021) 254:86

https://doi.org/10.1007/s00425-021-03740-y

REVIEW

Kopyor versus macapuno coconuts: are these two edible mutants

of Southeast Asia the same?
Adhityo Wicaksono1 · Reza Raihandhany2 · Jaime A. Teixeira da Silva3

Received: 8 July 2021 / Accepted: 21 September 2021

© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2021

Abstract
Main conclusion Kopyor and macapuno are two coconut mutants from Southeast Asia that are often described erro-
neously or interchangeably mismatched due to a lack of research, so future studies are encouraged.

Abstract Coconut (Cocos nucifera L.; Arecaceae), a widely distributed plant with popular culinary applications, especially
of the endosperm, has several nutritional and medicinal benefits. Two coconut mutants are widely recognized in Southeast
Asia, namely kopyor and macapuno, specifically in Indonesia and Philippines, respectively. Kopyor coconut is known for its
brittle solid endosperm while macapuno coconut is known for its gelatinous solid endosperm. Both mutant types have many
other synonyms in other countries. Over many decades, the biology of macapuno coconut, including endosperm anatomy,
histology, cytology, physiology, and genetics have been described, while kopyor coconut is still understudied. However,
some literature and websites erroneously describe kopyor as macapuno coconut, or consider them interchangeably, which is
an unintentional consequence of insufficient scientific research on these coconut mutants. Additionally, in Indonesia, there
is another local mutant in Banten called wax coconut (“kelapa lilin”) that some researchers claim as the actual Indonesian
macapuno coconut due to its strong resemblance to kopyor coconut. Unfortunately, wax coconut is not only understudied,
it is rarely documented. Additional evidence of their differences, in terms of morphological, biochemical and genetic
characteristics, is needed. Moreover, clear documentation will also be needed for a better comparison. Understanding the
differences between kopyor and macapuno coconuts will not only help to further clarify their scientific description in the
literature, but will also guide locals, researchers, and industries to characterize similar mutants, if found in specific regions,
for future study and bioprospecting.

Keywords Arecaceae · Elite coconut · Endosperm · Fruit · Palm

Introduction

Coconut (Cocos nucifera L.; Arecaceae) is a plant that origi‑

Communicated by Gerhard Leubner.
nated in Southeast Asia and is believed to be widely dis‑
* Adhityo Wicaksono tributed, by water, in Oceania, America, Africa, and other
adhityo.wicaksono@gmail.com parts of the world (Baudouin and Lebrun 2009). In terms
* Reza Raihandhany of global dispersion, coconut fruit is very resilient but pos‑
rezaraihan11@gmail.com sesses a loosely structured air layer, allowing it to float and
* Jaime A. Teixeira da Silva be dispersed by water (hydrochorous dispersal) to reach new
jaimetex@yahoo.com coastal destinations (Nilsson et al. 2010). Coconut trees are
broadly classified into tall and dwarf morphotypes (Perera
1
Division of Biotechnology, Genbinesia Foundation, Jl. et al. 2003). In terms of human use, coconut fruit is a popu‑
Swadaya Barat no. 4, Gresik Regency 61171, Indonesia
lar culinary ingredient and supplement (e.g. coconut oil,
2
Division of Botany, Genbinesia Foundation, Jl. Swadaya nata de coco, which is fermented from coconut water, etc.),
Barat no. 4, Gresik Regency 61171, Indonesia
the source of nutrition that possesses some ethnomedici‑
3
Independent Researcher, Ikenobe 3011‑2, P. O. Box 7, nal properties (e.g. detoxification medicine, anthelminthic,
Kagawa‑ken 761‑0799, Japan

13
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 86 Page 2 of 9
etc.), and even young coconut water or liquid endosperm is a
good component for plant tissue culture media as it contains
complex organic components, nutrients and plant growth
regulators such as auxins, cytokinins, and gibberellic acid
(Yong et al. 2009; Untari 2010; Dayrit and Nguyen 2020).
The known edible parts of the coconut are the apical
shoots, fruit endosperm and seedling cotyledons. Locally,
apical shoots or “umbut” in Javanese tradition, can be
cooked in local cuisines (Kristina and Hidayah 2019). Coco‑
nut has two types of endosperm, liquid and solid (Jirapong
et al. 2018), both of which are edible. The liquid endosperm
is actually a syncytial (cytoplasm with free nuclei) (Cut‑
ter et al. 1955 cit. Abraham and Mathew 1963), while the
solid endosperm is made up of cells with a range of chromo‑
somal ploidy numbers, including 3n, 6n, and 12n (wild type
being 2n = 32) (Abraham and Mathew 1963). On the micro‑
pyle side of the coconut, the embryo will grow inwardly
to develop a cotyledon (also referred to as a haustorium in
some articles) and outwardly to develop a shoot (plumule)
and roots (radicle) (Fig. 1) (Sugimura and Murakami 1990;
López-Villalobos et al. 2001; Pech et al. 2004). The coty‑
ledon (coconut is a monocotyledonous plant) is edible, has
a spongy, styrofoam-like texture, with a hint of sweetness
(Adhityo Wicaksono, personal experience), is locally called
“tombong”, “butuh”, or “kenthos” in some areas of Java
island, Indonesia (Azis and Lawan 2020; Heri Santoso, per‑
sonal communication), and could be considered as an equal
to endosperm as a food source. The coconut endosperm is
Fig. 1  Anatomy (a) and devel‑
opment of (normal) coconut
(a–c). Note the reduced level
of liquid endosperm indicating
absorption by the cotyledon
as the shoot grows and one of
three eyes is soft and func‑
tional, and serves as the embryo
growth opening, the rest are
plugged and impenetrable by
the embryo. Anatomical limited
by two dotted lines in a: Red—
seed, yellow—nut, and blue—
fruit. Illustration was inspired
from figures in the TNAU
Agritech Portal (https://​agrit​ech.​
tnau.​ac.​in/​horti​cultu​re/​horti_​
pcrops_​cocon​ut_​botany.​html)
13
Planta (2021) 254:86
normally edible as a dessert when it still young (6–7 months
old on the tree), with soft, mucous-like consistency, which
become more solid as the coconut matures (> 7 months old
on the tree), developing a tough endosperm (Gatchalian et al.
1994). Finally, coconuts are harvested for coconut milk and
oil (Sanful 2009; Agarwal and Bosco 2017). In addition to
than the association between different ages of coconut con‑
sumption and their distinct endosperm features, there are
also recorded cases of edible mutant (referred to as "elite"
in several papers) coconuts.
Mutant coconuts are very valuable due to their unique‑
ness, including their flavors. Examples are kopyor coconut,
macapuno (or makapuno) coconut, wax coconut (Indo‑
nesian: “kelapa lilin”), and canary coconut (Indonesian:
“kelapa kenari”). Canary coconut is an edible mutant coco‑
nut that originated from the Maluku province of Indonesia,
and is characterized by its crispy and brittle (like the canary
nut, Canarium indicum L., hence the origin of the name)
endosperm (Mashud et al. 2004). Kopyor coconut is also
an edible mutant coconut that is characterized by its loose,
velvety, soft, and brittle endosperm (Santoso et al. 1996),
and has many names: Dua Sap (Vietnam), Dikiri Pol (Sri
Lanka), Kopyor (Indonesia), Maphrao Kathi (Thailand),
Dahi Nariyel (Myanmar), Thairu Thengai (India), Dong
Kathy (Cambodia), and Niu Garuk (Papua New Guinea)
(Novarianto et al. 2014; Nguyen et al. 2016). In term of
nomenclature, kopyor is often mistakenly thought to be
similar to the macapuno coconut found in the Philippines.
Planta (2021) 254:86
The macapuno coconut, initially identified in the Philippines
(Gonzales 1914 cit. Nguyen et al. 2016), is another edible
mutant coconut that is characterized by its jelly-like or gelat‑
inous and thick endosperm (Lauzon 2005). In Indonesia,
the endosperm characteristics that have been described for
macapuno coconut are similar to those of wax coconut, in
addition to its appearance, which looks like melting wax
(Praktisi Humas 2016). This suggests that macapuno coco‑
nut is actually wax coconut rather than kopyor coconut. It
was also stated by Ismayanti (2013) that macapuno coconut
resembles wax coconut, and is characterized by its gelati‑
nous endosperm, rather than a brittle endosperm, which is
the characteristics of kopyor coconut). This view was rein‑
forced by a statement by Prof. Sudarsono (IPB University,
Indonesia; academic supervisor in Ismayanti (2013)) in an
interview, quoting “Macapuno coconut exists in Banten and
has been cultivated for generations, (there) it is called wax
coconut (“kelapa lilin”) because its endosperm resembles
the melting wax” (Putra 2016).
In this short review, characteristics of the morphology,
development, and physiology between kopyor and maca‑
puno coconuts, including the Indonesian wax coconut,
will be elaborately described based on the existing litera‑
ture. Based on the available evidence, we propose whether
kopyor, macapuno, and wax coconuts are the same or differ‑
ent mutants. We are also of the belief that there is a body of
literature that may have erroneously or mistakenly described
these mutants. This aspect will also be highlighted at the
end of this review to seek clarification regarding the naming
of these mutants and their characteristics, to offer guidance
for future research, and to serve as a cautionary note for
those wishing to cite such studies when publishing their own
papers. In addition, by providing a synthesized understand‑
ing of the differences between these mutant coconuts, future
prospects for conventional and biotechnological breeding
can be more realistically established.
Coconut endosperm
Biology and development
There are four types of endosperm development: (1) nuclear
type, in which the primary endosperm nucleus divides
within a single cytoplasm, forming a syncytial (multinu‑
cleated cell), ultimately resulting in cellularization (e.g.
Orontioideae and Aroideae) (Tobe and Kadokawa 2010);
(2) cellular type, in which the endosperm is formed by
cell division right from the beginning from the primary
endosperm nucleus (in Lemnoideae, Pothoideae, and
Aroideae) (Tobe and Kadokawa 2010); (3) helobial type, in
which cell division in the endosperm is transverse, resulting
in the independent development of micropylar and chalazal
Page 3 of 9 86
chambers (in Pothoideae, Monsteroideae, and Aroideae)
(Tobe and Kadokawa 2010); (4) ruminate type, where the
seed coat (testa) grows inward and dissects the endosperm
(in some members of the Arecaceae) (Zona 2005). Coco‑
nut endosperm adopts a combination of both nuclear and
cellular types of endosperm development, forming a mult‑
inucleated cell in a liquid form (liquid endosperm), after
which solid endosperm cells are formed next to the testa,
as cellularization occurs (Demason 1997). At an early stage
(up to 5–6 months of fruit development), the solid coconut
endosperm appears to be translucent, jelly-like, soft, and
can be peeled easily (Fig. 2c). At a later stage, the solid
endosperm becomes highly solidified, robust, and hard
to peel (Figs. 2d, 3a). In parallel development, the coco‑
nut embryo grows from the zygote, forming the coconut
seedling that consists of the cotyledon mass that absorbs
nutrients from the endosperm inside the seed, while the
plant grows outside the coconut fruit (Foale et al. 2020).
The normal coconut endosperm and embryo development
are described in Fig. 2e and also Fig. 3d. The cotyledon
will absorb the required nutrients for plumule and radicle
growth. The coconut shell (endocarp) is dense and robust,
but the embryo is located proximally to the one only func‑
tional stoma (known as the “eye”) out of three, where it is
softer than the remaining two “eyes”, allowing the embryo to
grow outwards by penetrating this spot (Hill 1937). In terms
of cytology, coconut has 32 diploid chromosomes (2n = 32)
(Teulat et al. 2000), while endosperm nuclei have triploid
(3n) nuclei, and fewer have hexaploid (6n) and dodecaploid
(12n) nuclei (Abraham and Mathew 1963).
Kopyor and macapuno
In the mutant kopyor coconut, the solid endosperm appears
to be malformed (Fig. 3b) the endosperm appears to be loose,
peeling off into the liquid endosperm by its own weight.
At an early stage (3–4 months old), the solid endosperm
appears to be less translucent than the normal coconut, and
still possess an ordered layer even though some parts have
already loosened (Fig. 2f). At a later stage (5–6 months old),
the solid endosperm becomes looser (Figs. 2g, 3b). Whereas
the embryo of a normal coconut develops as an orderly layer
of the solid endosperm (Fig. 3d), the embryo of the kopyor
coconut, due to its brittle and loose endosperm, becomes
exposed to the surface and to the liquid endosperm cavity
(Fig. 3e). While on the tree, the sound made by kopyor fruit
distinguishes it from normal coconut (Suppl. material S1;
young normal coconuts make no sound) when shaken by
hand, and has a clear liquid sound. Young kopyor coconuts,
when shaken by hand, produce a “light” solid sound inside
a liquid (Suppl. material S2), and will become a “heavier”
sound as more solid endosperm tissue peels off into the liq‑
uid (Suppl. material S3). The kopyor coconut is produced
13
 86 Page 4 of 9
Fig. 2  Endosperm development of normal coconut (a–d) versus
kopyor coconut (f–h). In the first 1–3 months, the young normal
coconut still has a tiny undeveloped seed cavity (a, b). The young
coconut (c for normal; f for kopyor), approximately 5–6 months old
on the trees, has a smooth, jelly-like solid endosperm (young normal
coconut), while the young kopyor coconut has both a normal-like
solid endosperm and a brittle solid endosperm. The mature coco‑
nut (d for normal; g for kopyor), approximately 7 months old on the
trees, has a solid and dense endosperm in the normal coconut, and an
irregular and brittle solid endosperm in the mature kopyor coconut.
Fig. 3  Comparison of the solid endosperm of a normal coconut (a),
a kopyor coconut (b), and a macapuno coconut (c). From the micro‑
pyle region of the coconut, the embryos (purple arrows) are exposed:
embryo of the normal coconut is sunken under the robust coconut
solid endosperm (d), embryo of the kopyor coconut is exposed due to
a brittle and loose solid endosperm (e), and embryo of the macapuno
coconut is sunken in the gelatinous solid endosperm (f). Notes: Blue
bidirectional pointers indicate proximal (towards the coconut core)
13
Planta (2021) 254:86
The liquid nuclear to cellular endosperm formation in a normal coco‑
nut (e) and in a hypothetical version of the kopyor coconut, where
the solid endosperm is malformed (h). Notes: Sliced young coconut
fruit (a, b) are discolored due to oxidation of their metabolites upon
cutting and exposure to air. Photo in d is the same image as Fig. 3a.
Blue lines, solid-textured endosperm; cyan lines, young jelly-like
endosperm; red lines, soft/gel-like endosperm; green lines, endocarp;
orange arrows, testa; purple arrows, primordial seed cavity. Coco‑
nut liquid nuclear endosperm formation was inspired from Tobe and
Kadokawa (2010). Scale bars: a, b 2.7 cm, c, d, f, g 1 cm
and distal (towards the coconut husk and skin) orientations. Photo
in a is the same image as Fig. 2d. Coconut anatomical layers: blue
lines, solid-textured endosperm; red lines, soft/gel-like endosperm;
green lines, endocarp; orange arrows, testa. Photo in c was photo‑
graphed by Enzo Pinga, and used with kind permission. The photo of
f was obtained from the Lunti Garden website (https://​lunti.​ph/​produ​
cts/​embryo-​cultu​red-​macap​uno-​mutant-​cocon​ut), and used with kind
permission. Scale bars = 1 cm
Planta (2021) 254:86 Page 5 of 9 86

only when the recessive endosperm mutant genotype is

Sisunandar et al. (2018); HEC composition is avail‑

formed (kkk) (Maskromo et al. 2016), hence kopyor fruit

27 ± 1 °C, then 26–29 °C; darkness for 4 weeks, then 14-h able in Sisunandar et al. (2018) supplementary
can only be found in an orchard on trees with the kopyor
recessive allele (k). Genetically and physiologically, it is still
unknown how the kopyor coconut is formed. However, it is
possible that solid endosperm formation is disrupted during

De Guzman and Del Rosario (1964)

development, leading to a loosened layer of solid endosperm
(Fig. 2h). The embryo of a kopyor coconut cannot grow to

AC activated charcoal, h hour(s), HEC hybrid embryo culture, IAA indole 3-acetic acid, IBA indole-3-butyric acid, mon month(s), PP photoperiod, RH relative humidity
the mature stage, and has to be grown via embryo rescue
(Table 1). As it grows into an adult coconut palm, the kopyor
coconut tree is classified as a dwarf coconut (Nguyen et al.

Warisno (1998)
2016).

References

materials
Visually, the macapuno coconut has an extra thick layer
of solid endosperm with a jelly-like appearance of the inner
layer (Fig. 3c). Unlike the kopyor coconut, the embryo of
the macapuno coconut is fully submerged into the solid
endosperm, although the inner part is in contact with the

­ −2 ­s−1; success rate from cul‑

24–26 °C; 60–70% RH; light: 1000 lx with 15-h PP; rou‑

­ −2 ­s−1, then mini growth chamber

ture growth, development, and acclimatization = 90%

jelly-like inner layer (Fig. 3f), thus giving the impression

In shaker with 75 rpm for 8–12 h/day for 6–8 weeks

that the embryo is growing normally. However, the maca‑

Embryo taken from fruit 11 mon after pollination;

puno coconut embryo also cannot reach the mature stage
due to malnutrition. According to Nguyen et al. (2016)
(Fig. 4), the reserve glucomannans will be enzymatically

25 °C; dark for 3 weeks, then in light

processed into fructose, mannose, and galactose, finally

tinely subcultured for 8–12 weeks

providing nourishment for the embryo that later will be

with 14-h PP at 40 µmol m

absorbed by the cotyledon, allowing the coconut to sprout.
In macapuno coconut, the activity of β-mannase increases,

PP at 25 ± 2 µmol m
leading to accumulation of mannobiose, mannotriose, and
Environment details

galactomannobiose in the endosperm. However, the maca‑

puno coconut lacks converting enzymes, α-D-galactosidase
Table 1  Embryo rescue protocols for kopyor and macapuno (based on Biddle et al. 2020)

and β-mannosidase, leading to the absence of nutrition in the

form of mannose and galactose for the embryo, thus embryo
growth is inhibited. In a normal coconut, tryptophan (amino
acid) will be processed into indole-3-acetic acid (IAA) or
Liquid embryo culture: White modified de Guzman, White

auxin inside the endosperm, but the pathway is regulated by Root induction and acclimatization: Mini growth chamber
modified Norstog, and White modified Eeuwens media

Growth: HEC (in light), Rillo et al. (2002) vitamins, and

with aerated HEC (in light), 1 µM IBA, no vitamins or

Solid culture for seedling: Murashige and Skoog (1962)
with 2% dextrose, pH 6.0 (before autoclaving), shaker

basal media with 6% dextrose, 10 mg/l IAA, 0.5 mg/l

For growth and plant development: White (1964), 25%

the activity of peroxidase and IAA-oxidase. In macapuno,
sucrose, cocopeat: rice charcoal substrate (1:1, v/v)
Germination: HEC, Rillo et al. (2002) vitamins, 6%
75 rpm, 8–12 h of shaking per day for 6–8 weeks

IAA biosynthesis is upregulated because peroxidase and

IAA-oxidase are lacking. These observations were extrapo‑
lated from Nguyen et al. (2016). Due to inhibited growth,
filtered coconut water, and 1.2% agar

the macapuno embryo, just like in the kopyor coconut, has to

be grown via embryo rescue (Table 1). Just like the kopyor
coconut, the macapuno coconut carries a recessive allele
(m), thus indicating that fruit with a macapuno endosperm
has only recessive alleles (mmm). Naturally, it has to grow
sucrose, and 0.2% AC

in a coconut orchard laden with macapuno-bearing trees to

IBA, and 5% AC

intensify the probability of occurrence of this mutant. How‑

3% sucrose

ever, unlike kopyor, which is classified as a dwarf coconut

palm, macapuno is classified as a tall coconut palm/tree
Cultured plant Media

(Nguyen et al. 2016).

Cytologically, the macapuno coconut endosperm has
been observed to have an irregular cell arrangement with‑
out cell adhesion, and cell morphology is irregular and
Macapuno
Kopyor

vary in size but larger than cells of the normal coconut, and
cells divide into multinucleate cells by budding or cellular

13
 86 Page 6 of 9
Fig. 4  Physio-developmental pathway in a normal coconut, where
sugars are produced to feed the embryo and indole-3-acetic acid
(IAA) is constantly regulated (a) and a macapuno/wax coconut,
where some sugars are upregulated but having only fructose as the
final product due to the lack of some enzymes, leading to a malnour‑
ished embryo, and the accumulation of IAA (b). Notes: dark green,
normal enzymatic and developmental processes; blue, regulative
pathway; red, downregulated/reduced pathway and inhibited growth;
magenta, upregulated/increased pathway and altered growth; nor‑
mal arrows, normal progression pathways; dotted arrows, inhibition
effect; matchstick lines, progressive regulation pathways. Schematics
were inspired from Nguyen et al. (2016)
“fragmentation” rather than normal cytokinesis (Abraham
et al. 1965; Ramirez 1986). Additionally, the endosperm
cells possess much larger ploidy numbers than in normal
coconuts at the mature stage of development (i.e., in maca‑
puno coconut, cells with ploidy numbers of 3n, 6n, 9n, 12n,
24n, 48n, and beyond, were found [3n = 44.90–78.44%,
6n = 14.00–41.56%, 9n = 0.22–0.89%, 12n = 0.44–5.78%,
24n = 0.00–1.55%, and 48n and beyond = 0.00–4.65%,
relative to 2n = 0.00–0.89%; observed in 1800 endosperm
cells, with 450 cells per bunch of fruit, and with bunches 6,
7, 8, and 9 counted from the top of the inflorescence]) as a
result of c-mitotic division (aberrant mitosis at metaphase
stage) or nuclear fusion after nuclear division is complete
in aberrant mitosis (Abraham and Mathew 1963; Abraham
et al. 1965; Ramirez 1986). No in-depth study has been
conducted on kopyor coconut endosperm to understand its
cytology and if there are chromosomal changes.
13
Planta (2021) 254:86
Possible mismatched information
in publications
In some texts, kopyor coconut has inaccurately been used
interchangeably as macapuno coconut despite their differ‑
ences in endosperm anatomy, as described above. Some
reports and journals (Uswadi et al. 1979; Rival 1991;
Sukendah et al. 2011; Novrianto et al. 2014) mistranslated
kopyor as macapuno. One paper (Sukendah 2018) mistook
the description of the physiological defect of macapuno
(citing Mujer et al. 1984; a paper on macapuno coconut)
as being of kopyor coconut. Another study (Sisunandar
et al. 2018) describes kopyor coconut as a result of spon‑
taneous mutation, citing Nguyen et al. (2016), who were
describing macapuno coconut. Another paper (Antu et al.
2021) noted that kopyor is referred to as macapuno in the
Philippines. One magazine also mentioned the same claim
that kopyor is referred to as macapuno coconut in the Phil‑
ippines (Winarno 1981), suggesting that this mismatch has
occurred from at least the 1980s.
When searching for the terms “macapuno kopyor coco‑
nut” and “kelapa makapuno kopyor” on Google, there are
many other confusions, interchangeable use of both forms,
and mistranslations, even in English, Indonesian and Java‑
nese Wikipedia (macapuno: https://​en.​wikip​edia.​org/​wiki/​
Macap​uno; kopyor: https://​id.​wikip​edia.​org/​wiki/​Kelapa_​
kopyor; https://​j v.​wikip​e dia.​o rg/​wiki/​Kramb​i l_​kopyor),
while in Vietnamese Wikipedia, dua sap is shown with
the image of a macapuno coconut: https://​vi.​wikip​edia.​
org/​wiki/​Dừa_​sáp. Even on the English Wikipedia page,
“macapuno” erroneously mixes references about kopyor
(Santoso et al. 1996; Sukendah and Sudarsono 2009) and
macapuno (Zuñiga 1953; Rillo 1999; Areza-Ubaldo et al.
2003; Angeles et al. 2018). Additionally, at least 37 web‑
sites, including wiki’s, news article, blogs, photo galleries,
social media, and even governmental pages have described
kopyor coconut as macapuno coconut (Supp. Table 1).
This lax differentiation between both forms of coconut
should encourage better comparative studies between
macapuno and kopyor coconuts. Additionally, since wax
coconut (“kelapa lilin”) is claimed to be the real maca‑
puno (according to Prof. Sudarsono in Praktisi Humas
2016; and Putra 2016), no online documentation is avail‑
able, so scientific studies dedicated to the conservation,
characterization, and documentation of wax coconut are
needed. To date, only one confirmed observation of wax
coconut appears to exist. It was found, following exten‑
sive searches, on Instagram. The authors communicated
with the account holder, Silviana Salim from Banten, who
confirmed that the wax coconut was bought from a local
coconut vendor in November 2018 at Tanjung Kait coastal
area, Tangerang district, Banten, Indonesia). Indeed, that
Planta (2021) 254:86
image reveals visual similarities in the endosperm of wax
(Fig. 5) and macapuno (Fig. 3c, f) coconuts. Wax coco‑
nut is also known as a local delicacy by some in Banten
(Java Island, Indonesia), although some people dispose
the coconut since its appearance is less appetizing and
resembles a gooey stale coconut due to missing informa‑
tion between individuals who knows this fact and others
who do not (Tina Handimuljana, personal communica‑
tion). Hence, future studies are also encouraged to allow
for an understanding of the nutritional value of wax or
macapuno coconut in Indonesia, complemented by other
research on this germplasm.
Conclusion
Kopyor and macapuno/makapuno are mutant coconuts.
Kopyor coconut has a brittle endosperm that is hypo‑
thetically deformed during early formation of the solid
endosperm. Macapuno coconut, on the other hand, has
gelatinous endosperm, a solid endosperm that is rich in
galactomannan, and a liquid endosperm that is rich in IAA,
both caused by the lack of enzymatic production. Maca‑
puno coconut also has disrupted cytokinesis in endosperm
cells, including polyploid cells. Both macapuno and kopyor
coconuts are considered as delicacies, especially in Indone‑
sia (kopyor) and the Philippines (macapuno). In Indonesia,
kopyor coconut biology is still understudied, leading to some
confusion and mistranslaton as macapuno coconut in some
papers and media. The real macapuno coconut in Indonesia
is claimed to be the wax coconut or “kelapa lilin”. However,
wax coconut is scarcely studied and no documentation is
available, so a detailed comparison between wax, macapuno,
and kopyor coconuts is necessary.
Fig. 5  Local documentation of wax coconut from Banten, Indonesia,
revealing endosperm visual similarities with macapuno (Fig. 3c, f).
Photo taken and provided by Silviana Salim, with kind permission
Page 7 of 9 86
Author contribution statement All authors contributed
equally from ideation to writing and editing of the paper.
Supplementary Information The online version contains supplemen‑
tary material available at https://​doi.​org/​10.​1007/​s00425-​021-​03740-y.
Acknowledgements We thank Enzo Pinga from the Philippines for
providing the picture of macapuno, and Jalexander Hongco for his per‑
mission to use the picture from Lunti Garden Philippines website (lunti.
ph). We are grateful to Ibu Helena Sutisna (Rumah Kopyor, Bandung,
Indonesia) for discussion and help to find the wax coconut (despite its
rarity), and for selling us the kopyor coconuts, which is considered
to be quite an uncommon commodity to find. We are grateful to Ibu
Silviana Salim from Banten, Indonesia, for providing the picture of
wax coconut and some background information regarding the location
where it was found. We are also grateful to Ibu Tina Handimuljana
(Rumah Tjoen, Banten, Indonesia) for discussion and her generosity
for sharing information (from herself and gathered from locals) about
the wax coconut and for her patience and willingness to find us a wax
coconut (although, after 3 months, no wax coconuts were found on her
coconut farm). Finally, we thank the director of the Genbinesia Foun‑
dation, Heri Santoso, for sharing his ethnobotanical understanding of
some of the observations made in this paper.
Data availability statement All data generated or analyzed during
this study are included in this published article (and its supplementary
information files).
Declarations
Conflict of interest All authors declared no conflicts of interest.
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