e-ISSN 2477-9148
11
Elaeis oleifera: a neglected palm from Ecuador
Montúfar et al.
REVISTA ECUATORIANA DE MEDICINA Y CIENCIAS BIOLOGICAS
Volumen 39. No. 1, Mayo 2018
Elaeis oleifera (Kunth) Cortés: A neglected palm
from the Ecuadorian Amazon
Elaeis oleifera (Kunth) Cortés: una palma olvidada
de la Amazonía ecuatoriana
Rommel Montúfar
1
*, Claude Louise
2
, Timothy John Tranbarger
1,3
1
* Ecology and Genetics Laboratory, Ponticia Universidad Católica del Ecuador (PUCE), Ecuador. (rjmontufar@
puce.edu.ec)
2
Palmelit sas Parc Agropolis, 2214 Boulevard de la Lironde, Montferrier-sur-Lez, 34980, France. (claude.louise@
palmelit.com)
3
UMR DIADE, Institut de Recherche pour le Développement, Université de Montpellier, Montpellier, France.
(timothy.tranbarger@ird.fr)
*Corresponding author e-mail: rjmontufar@puce.edu.ec
doi.org/10.26807/remcb.v39i1.584
Recibido 02-03-2018 ; Aceptado 23-03-2018
ABSTRACT.- Ecuador has an outstanding diversity of palm species, some of which have been well studied, certain others remain
an enigma, however. A particular case is the American oil palm, Elaeis oleifera (Kunth) Cortés, rst recorded in Ecuador in 1986.
The genus Elaeis has a trans-Atlantic (Africa-America) distribution, with E. oleifera from the Neotropics, and E. guineensis Jacq.
from Africa. It has been hypothesized that E. oleifera derives from populations of E. guineensis, which diverged 15 million years
ago. At the local level, the populations of E. oleifera have a disjunct distribution, with isolated populations in Central America,
the Amazonia basin, the Guianas, Chocó and the Caribbean Cost of Colombia and Venezuela, frequently associated with human or
archaeological settlements. Despite the spatial and historical separation between the two species, there are no reproductive barriers
to the generation of fertile hybrids. This important reproductive characteristic has allowed E. oleifera to become a major source
of genetic variation for the improvement and adaptability of commercial populations of E. guineensis throughout the tropics. The
Ecuadorian populations of E. oleifera from Taisha, with morphological, reproductive and agronomically important biochemical
characteristics, have been used for the creation of commercial hybrids, which today are planted in many tropical regions.
KEYWORDS: Ecuador, Elaeis guineensis, fruit abscission, oil palm, palms.
RESUMEN. - El conocimiento sobre la biología de las palmas ecuatorianas es todavía limitado. Un caso particular constituye la
palmera aceitera americana, Elaeis oleifera (Kunth) Cortés descrita para Ecuador en el año 1986. El género Elaeis tiene una distri-
bución trans-Atlántica (África-América), con E. oleifera proveniente del Neotrópico y E. guineensis Jacq. del África. Se ha sugeri-
do como hipótesis que E. oleifera deriva de poblaciones de E. guineensis, las cuales divergieron hace 15 millones de años. A nivel
local, las poblaciones de E. oleifera presentan una distribución disyunta, con poblaciones aisladas a lo largo de la América Central,
Amazonía, Guyanas, el Chocó y la costa caribeña de Colombia y Venezuela. Están frecuentemente asociadas a asentamientos hu-
manos o arqueológicos. A pesar de la separación espacial e histórica entre ambas especies, no existen barreras reproductivas para
la generación de híbridos fértiles. Esta importante característica reproductiva ha permitido que E. oleifera constituya actualmente
la principal fuente de información genética para el mejoramiento y adaptabilidad de las poblaciones comerciales de E. guineensis
a lo largo del planeta. Las poblaciones ecuatorianas de E. oleifera provenientes de Taisha (Provincia de Morona Santiago) –con
características morfológicas, reproductivas y bioquímicas agronómicamente importantes– han sido utilizadas para la creación de
híbridos comerciales que, actualmente, se siembran en las regiones tropicales.
PALABRAS CLAVES: abscisión del fruto, Ecuador, palma aceitera, Elaeis guineensis.
Artículo de revisión
12
REMCB 39 (1): 11-18, 2018
INTRODUCTION
Ecuador harbors one of the highest palm species di-
versities in the world, with more than 136 species
(Valencia and Montúfar 2013). In spite of this extraor-
dinary diversity, and the multitude of uses of many
palm species, much knowledge remains to be acquired
about the natural history. One example is the little-stu-
died species Elaeis oleifera (Kunth) Cortés (e.g. the
American oil palm) which has a high potential for the olea-
ginous industry. The objective of this paper is to review
the literature available on the Ecuadorian populations of
E. oleifera. As information is limited, this paper
represents progress towards increasing our knowledge
about the natural history, distribution and characteristics
of this intriguing yet poorly studied tropical native
Ecuadorian palm.
The genus Elaeis.- The genus Elaeis (subfamily
Cocoseae, subtribe Elaeidinae; Dranseld et al. 2008)
of the palm family (Arecaceae) is present in the tropical
regions of Africa and in Central and South America.
Elaeis is a small palm genus comprised of two species:
E. oleifera, from the Americas, and its sister species
Elaeis guineensis Jacq., from Africa, which us commonly
known as the African oil palm. Elaeis guineensis is
widely cultivated throughout the tropical regions for its
fruit, which yields palm oil, and is globally considered
the most important source of edible vegetable oil in
both production and trade, accounting for one-third of
worldwide vegetable oil production in 2009 (Murphy
2014; Vijay et al. 2016). In recent decades, scientic
interest has led to research being focused on the
physiology, genetics and genomics of E. guineensis; in
particular the agronomic characteristics related to yield,
and the remarkable capacity to synthesize and store
lipids in both the fruit mesocarp (palm oil) and the kernel
(kernel oil) tissues (Murphy 2006; Murphy 2009; Bourgis
et al. 2011; Tranbarger et al. 2011; Dussert et al. 2013;
Singh et al. 2013a; b; Corley and Tinker 2016; Guerin
et al. 2016). Furthermore, there is a high quality draft
of the E. guineensis genome, and “omic” technologies,
bioinformatics, marker assisted selection (MAS) and
transgenic technologies have been and continue to be
developed in order to accelerate genetic improvements
(Singh et al. 2013; Murphy 2014).
The biogeography of the Elaeis genus is still a puzzle.
Most palm genera are strictly endemic and have
evolved within the context of a specic continental
area (Henderson et al. 1995); however, the genus Elaeis
displays a trans-Atlantic disjunction given its presence
on the African and American continents (Renner
2004). It has been inferred that American E. oleifera
populations are derived from ancient Elaeis populations
which dispersed from Africa via sea currents before
the end of the Miocene (Renner 2004; Dranseld et al.
2008). Based on two dierent estimates, one on a dated
molecular phylogeny of the palm family, and the other
on Elaeis genome sequences, these species diverged
from 15–20, or 51 million years ago, respectively (Baker
and Couvreur 2013; Singh et al. 2013). The species that
we now know as Elaeis oleifera was originally described
with collections from Cartagena, Colombia, as Alfonsia
oleifera by Kunth (1815). It was later transferred to Elaeis
by Cortés as Elaeis oleifera (Kunth) Cortés (1897), and
still later it was recombined as Corozo oleifera (Kunth)
Bailey (1933). In parallel, another species was described
from Brasil, Elaeis melanococca Mart. (1824), but that
name is now treated as a synonym of Elaeis oleifera.
Elaeis oleifera has disjunct populations located in
Central America (Honduras to Panama), the Amazonia
basin, the Guianas, the Chocó and the Caribbean coast of
Colombia and Venezuela, growing naturally in the
tropical forest at 0–500 meters above sea level (masl),
with optimal temperatures of 23–30 ºC and an optimal
annual rainfall of 1400–2500 mm (Ecocrop 2007).
Despite their evolutionary distance, E. oleifera can be
crossed with E. guineensis to form interspecic hybrids
(O x G hybrids) which can be fertile (Singh et al. 2013b;
Corley and Tinker 2016). E. oleifera is thus used as
the female parent source for genetic variation that can
be useful in breeding programs that target the creation
of improved commercial varieties of E. guineensis
(Barcelos et al. 2015).
At the morphological level, while the two species are
fairly similar in general appearance, there are clear di-
erences in their vegetative and reproductive structures,
between E. oleifera and E. guineensis which reect their
trans-Atlantic disjunction (Corley and Tinker 2016; Ta-
ble 1). Firstly, E. oleifera is shorter, which may be the
result of a slower annual height increase as compared
with E. guineensis, and E. oleifera can display procum-
bent trunk growth. While root development is similar to
E. guineensis, adventitious roots can develop along the
entire length of the procumbent trunk. The leaf structure
of E. oleifera is markedly dierent from E. guineensis, in
that the leaets of E. oleifera lie in a single plane, while
the leaets of E. guineensis are arranged in groups and
project in dierent planes. Another striking dierence is
the brous spathe that covers the female inorescence of
E. oleifera, which remains until the fruit have ripened.
Elaeis oleifera fruit are smaller and are often found to be
parthenocarpic, while the fruit bunches typically display
a conical shape, pointed at the top (Draneld et al. 2008;
Balslev 1987; Corley and Tinker 2016). Elaeis oleifera
is often found in damp, swampy areas, near riverbanks
or in pastureland. It is shade tolerant in comparison with
13
REVISTA ECUATORIANA DE MEDICINA Y CIENCIAS BIOLOGICAS
Elaeis oleifera: a neglected palm from Ecuador
Montúfar et al.
E. guineensis, which can be considered a pioneer species
and is intolerant of shade. A more detailed list of the re-
ported dierences between E. oleifera and E. guineensis
is found in Table 1.
Elaeis in Ecuador,- The history of Elaeis in Ecuador is
heavily inuenced by the introduction of the African oil
palm, E. guineensis, as a crop for oil production. The rst
reported introduction of Elaeis guineensis to Ecuador
was in 1940 at the tropical experimental station of the
Instituto Nacional de Investigaciones Agropecuarias
(INIAP, Los Ríos Province) in Pichilingue; and in 1959
it was introduced to the Estación Agroforestal de San
Lorenzo (Esmeraldas Province, Acosta-Solís 1971).
Elaeis guineensis is currently cultivated in four main
areas in Ecuador: Quinindé (Esmeraldas), San Lorenzo
(Esmeraldas), the Southern Coast (El Oro, Guayas) and
the Amazonia (Estimations of Asociación Nacional de
Cultivadores de Palma Africana del Ecuador, ANCUPA
in 2005 and in Danec S.A. 2018). According to current
estimates, oil palm plantations constitute approximately
300 kha of total area harvested in the four zones,
making Ecuador the third largest producer (following
Colombia and Guatemala) of palm and kernel oil in
the Americas, with 620 000 MT produced in 2016
(FAOSTAT 2018; IndexMundi 2018; United States
Department of Agriculture - USDA 2018; Table 2).
However, as elsewhere in the tropics, the development of
the Ecuadorian oil palm industry has been controversial;
mainly due to the conversion of tropical rain forest
into areas for oil palm cultivation, resulting in a loss
of biodiversity along with potential negative socio-
economic consequences (Vijay et al. 2016; Marin-
Burgos and Clancy 2017).
By 1965, a 39 ha experimental plantation of E. oleifera
—apparently planted with seeds imported from outside
of Ecuador— was established at the experimental INIAP
(Instituto Nacional de Investigaciones Agropecuarias)
station in Santo Domingo de los Tsáchilas (Borgtoft Pe-
dersen and Balslev 1993). The rst botanical report of
native E. oleifera in Ecuador dates from 1986 (Balslev
and Henderson 1986; Balslev 1987). The botanists Hen-
rik Balslev (Aarhus University, Denmark) and Andrew
Henderson (New York Botanical Garden) described the
rst native E. oleifera population of 10 individuals in the
locality of Taisha (450 masl) in the province of Morona
Santiago (Ecuadorian Amazon). Later, wild E. oleifera
populations were also discovered in the Pastaza (Figure
1; AAU Herbarium Database 1990, 2011) and Orellana
provinces (Nuevo Rocafuerte, Barba et al. 2014). The
Danec S.A. Company, with French partners CIRAD/Pal-
mElit SAS, conducted their own prospection in 2003 in
the Taisha area and planted sample populations of this E.
oleifera-Taisha material in Quinindé and in Shushundi,
Sucumbíos (Figure 2). INIAP maintains a seed bank co-
Table 1. Morphological differences between Elaeis oleifera (Kunth) Cortés and E. guineensis.
Characteristic
E. oleifera
E. guineensis
Habitat
-damp or swampy areas; near or on the riverbank; pasture land; tolerant of shade
-disturbed areas; can act as a pioneer species; difficult to assess natural
habitat because often associated with human habitation; intolerant of
shade
Dispersal vector
-humans, undocumented
-animals (including humans) some bird species
Stem
-short and prostrate; only the terminal portion with the leaves is erect.
-1-6m tall, 40 cm diameter
(trunks lying on the soil for a distance of over 3-7 m)
-erect
-10 m tall, 30-50 cm diameter
Leaf /blade
-20-50 leaves
-sheath 20-40 cm long
-petiole 1.5-3 m long covered by short and thick spines on margins, no basal
swellings on the petiole
-rachis 2.9-6 m long
-blade 3 to 6 m long and 2.5 m wide
-40-60 leaves
-sheath 5 m long
-petiole > 1.2 m long covered by short lateral spines
-rachis c.a. 8 m
Pinnae (leaflets)
-30-90 linear and acuminate pinnae per side, middle ones up to 1 (1.2) m long, 4-6
cm wide
-regularly distributed along rachis and spreading in one plane
- >100 linear and acuminate pinnae per side, middle up to 1.3 m long, 6
cm wide
-arranged in groups (2 to 4) and spreading in different planes
Roots
-roots develop along whole length of procumbent trunk
-no procumbent trunk growth
Fruit
-ellipsoid-oblong, 2.5-3 cm long, 1.8-2 cm diam.
-weight range from 1.7 5.0 g in Colombia, 5-13 g in Brazil
-yellowish orange to red.
-prominent apical stigmatic residue
-commonly parthenocarpic fruit (up to 90%)
-spherical to ovoid, 2-5 cm long
- weight range from 6 to 20 g
-red to black
Fruit Bunch
-conical bunches 8 to 12 (30) kg
-max. number of fertile fruits reported was 5000
-ovoid bunches up to 100kg
-500 to 4000 fruit per bunch reported
Male inflorescence
-100 to 200 spikelets, 5 to 15 cm
-smaller with shorter anthers
-long peduncle
-long, finger-like, cilindrical spikelets, 10-20 cm lenght
Female inflorescence
-persistent spathe
-long penducle
-flowers sunk in the body of the spikelet
-period of anthesis is erratic and may last for 3 or 4 weeks or have two peaks
-long peduncle
-flowers subtended by a long bract
-period of anthesis in 36 to 48 hours to 1 week.
Pollinator
-Grasidius hybridus O’Brien and Beserra
-Elaeidobius kamerunicus Faust
Pollen
-elliptical shaped
-triangular shaped
Abscission Zone
-simple, multiple abscission zone cell layers with aligned nuclei not observed
-vascular tissue appears to be lignified in abscission zone.
-complex, multiple abscission zone cell layers characterized by aligned
nuclei
-vascular tissue not lignified in abscission zone.
Compiled from the following references: Zeven 1964; Borgtoft Pedersen and Balslev 1993; Henderson 1995; Smith 2015; Corley and Tinker 2016; Fooyontphanich et al. 2016; Auffray et al. 2017;
http://www.fao.orsȀ
Table 1. Morphological dierences between Elaeis oleifera (Kunth) Cortés and E. guineensis.
14
REMCB 39 (1): 11-18, 2018
llection of E. oleifera from dierent Ecuadorian provin-
ces, but very little information about this collection has
been made public (Ortega Cedillo et al. 2016). Beyond
these initial reports, no comprehensive description of the
wild E. oleifera populations present in Ecuador has been
made. Thus far, individual samples in Ecuador have been
reported in the eastern lowlands, mainly in the Morona
Santiago (Taisha, 450 masl) and Pastaza (500 masl) pro-
vinces, on poorly drained soil and in uvial plains (Bor-
chsenius et al. 1998). The disjointed local distribution
of E. oleifera in the Amazon has been explained by the
anthropogenic inuence on the palm’s dispersal (Balée
1989).
Genetic and phenotypic diversity of Elaeis in
Ecuador.- Several studies have used genetic markers
to describe the genetic and phenotypic diversity of
E. oleifera; however, these studies have examined
only (i) seed accessions from regional gene banks, or
(ii) individuals cultivated ex situ. To the best of our
knowledge, no studies of natural populations in situ have
been reported. Nevertheless, a study of ex situ individuals
from dierent countries with Simple Sequence Repeats
(SSRs) markers revealed that E. oleifera from Taisha
(Morona Santiago) are, surprisingly, more genetically
similar to populations from Pacic Colombia (Valley
of Sinú, Antioquia) than the geographically adjacent
populations from the Peruvian Amazonia ( Arias et al.
2015). At the phenotypic level, while yield components
(FFB-fresh fruit bunches, kg per palm year
−1
; BN-number
of bunches, palm year
−1
; MBW-mean bunch weight,
kg) were lower for Taisha individuals, other unique
qualitative traits were dierent in comparison with
populations from Colombia, Peru, and Brazil, including:
(i) 14–21% higher Mesocarp/Fruit (M/F) ratio (63%);
(ii) higher Fruit/Bunch (F/B) ratio for parthenocarpic
Table 2. Estimation of cultivated area of E. guineensis and E. oleifera x E. guineensis hybrids in Ecuador. Estimations
from Asociación Nacional de Cultivadores de Palma Aceitera del Ecuador ANCUPA and Danec S.A. in 2018.
Zone
E. guineensis
(ha)
Hybrids Total
(ha)
Hybrids Coari
1
(ha)
Hybrids Taisha
2
(ha)
Hybrids others
(ha)
Quinindé
203 500
6 500
5 000
1 000
500
San Lorenzo
2 500
22 500
19 500
2 500
500
Southern Coast
30 000
0
Amazonia
15 000
15 000
11 800
3 000
200
TOTAL
251 000
44 000
36 300
6 500
1 200
1
Coarí is an E. oleifera accession from Brazil.
2
Taisha is an E. oleifera accession from Ecuador.
Figure 1 Wild population of E. oleifera from Pastaza Province, Ecuador. (A) An adult individual in the primary forest; (B) Male
inorescence at early stage of development; (C) Spines along of the base of the petiole. Source: Henrik Balslev (#Balslev, H. 8512),
AAU Herbarium Data Base.
15
REVISTA ECUATORIANA DE MEDICINA Y CIENCIAS BIOLOGICAS
Elaeis oleifera: a neglected palm from Ecuador
Montúfar et al.
fruit; (iii) the absence of peduncular bracts; (iv) green
immature fruits; and (v) longer spikelet stalks (Arias
et al. 2015). In Ecuador, the genetic diversity of 40
individuals from the germplasm bank of INIAP in Santo
Domingo was analyzed with microsatellites markers,
and a low endogamy and high genetic variability was
shown (Ortega Cedillo et al. 2016).
Pollination of E. oleifera.- While there have been
a number of studies on the pollination process of E.
guineensis, the pollination of E. oleifera is poorly
understood (Corley and Tinker 2016; Auray et al.
2017). Evidence shows that the genus Elaeidobius of the
Coleoptera order, in particular Elaeidobius kamerunicus,
is the natural pollinator of E. guineensis. For E. oleifera,
the derelomine weevil, Grasidius hybridus (Coleoptera:
Curculionidae) was collected in a natural population of
E. oleifera in Taisha and is apparently a natural pollinator
of this species (Auray et al. 2017). In addition,
G. hybridus was reported as a crepuscular pollinator,
while E. kamerunicus was actively present in the
morning on ex situ populations of E. oleifera-Taisha
cultivated in the Ecuadorean Amazonia. In South
America, O x G hybrids must be pollinated manually
with E. guineensis pollen, due to the poor viability of
hybrid pollen, and the hybrid inorescences are less
attractive to E. kamerunicus. Therefore, research on the
pollination process of E. oleifera and O x G hybrids is
an important area of study (Meléndez and Ponce 2016).
Ethnobotanical Uses of E. oleifera.- In Ecuador, the
only indigenous name reported in the literature comes
from the Achuar communities, where it is known as
Yunchik (Borchsenius et al. 1998). No ethnobotani-
cal uses have been formally reported from Ecuadorean
populations of E. oleifera. However, there is a limited
amount of information from other countries about tradi-
tional uses, which include folk remedies, beverages, in-
sect-repellents and cooking (Smith 2015). Interestingly,
both E. oleifera and E. guineensis are typically closely
associated with human settlements and movement (Smi-
th 2015).
Genetic breeding programs and E. oleifera traits of
interest.- The introgression of genetic information from
E. oleifera to the widely cultivated E. guineensis through
the creation of O x G hybrids is a major objective of the oil
palm industry, mainly due to the resistance of E. oleifera
to lethal bud rot or fatal yellowing disease (Pudrición
del Cogollo, PC; Corley and Tinker 2016; Barcelos et al.
2015). Oil palm breeders have been successful in selecting
O x G hybrids with tolerance to lethal bud rot, and which
have yield components comparable to the average of E.
guineensis crosses. In addition, there is much interest in
transferring the higher oleic acid content of E. oleifera to
E. guineensis. Recently, a study with E. oleifera-Taisha
showed the results of an 8-year evaluation of O x G
hybrids (E. oleifera-Taisha x E. guineensis-Avros). This
study revealed the outstanding potential of Ecuadorean
populations of E. oleifera from Taisha. For example,
some of these O x G hybrids showed tolerance to PC and
to other diseases, had low annual growth rates, uniform
anthesis, very few spathes that cover the inorescence,
and a long peduncle that makes for easier harvest (Barba
and Baquero 2013). Additionally, oil derived from the
fruits of hybrids had a higher concentration of oleic
acid, which is attractive to the vegetable oil industry. In
another article it was found that E. oleifera-Taisha and
an intraspecic E. oleifera hybrid “Manaos/Taisha” had
total fruit weights in the range of the E. guineensis (Lieb
et al. 2017).
A recent study found E. oleifera-Taisha individuals
planted ex situ in Quinindé, (Ecuador) do not drop their
fruit from the bunch, which normally occurs naturally
through the fruit abscission process (Fooyontphanich et
al. 2016). In addition, the abscission zone of E. oleifera
was markedly dierent from that of E. guineensis
(Table 1). Fruit abscission is an important agronomic
characteristic whose control is of interest to oil palm
breeders in order to facilitate harvest and reduce the
impact of oil acidication due to lipase activity induced
in damaged abscised fruit (Morcillo et al. 2013). A recent
survey of the E. oleifera-Taisha population in Quinindé
conrmed the non-shedding character of certain
individuals. In particular, one individual was found
to have seedlings that develop from fruit still attached
to the bunch (Figure 2E). Furthermore, an in vitro
phenotype test of abscission revealed that no separation
in the abscission zone took place after a 24-hour test
period (Figure 2F; Fooyontphanich et al. 2016). This
E. oleifera-Taisha individual provides genetic material
important for understanding the abscission process in
owering plants, in addition to the possible introgression
of genetic information to modify the abscission process
in E. guineensis.
These limited examples show the importance of des-
cribing and conserving the local biodiversity of
E. oleifera-Taisha, which is represented by a single lo-
cality in the Ecuadorian Amazonia. A more complete
exploration of the Ecuadorian diversity of E. oleifera
and the implementation of a national program to protect
these natural populations as a source for traits and genes
that could be benecial to a sustainable oil palm industry
is thus of great importance.
Conservation of E. oleifera.- Elaeis oleifera was not
assessed for the International Union for Conservation of
Nature (UICN) red list of Ecuador (Montúfar et al. 2011)
16
REMCB 39 (1): 11-18, 2018
because this inventory was focused on endemic species.
However, due to its limited known distribution, the few
botanical collections reported, and its economic impor-
tance as a source of genetic material for the oleaginous
industry, this species could be considered endangered
and should therefore be included into the National Agen-
da of Research of Biodiversity (INABIO 2017).
In conclusion, the cultivation of oil palm in industrial
plantations is controversial in tropical countries, inclu-
ding in Ecuador. However, it is clear one objective, and
a major worldwide challenge, is how to develop more
productive, sustainable genetic material and cultivation
practices that will reduce the pressure on native tropi-
cal rain forests. E. oleifera clearly could have a positive
impact on both biodiversity conservation and the gene-
tic improvement of the oil palm. In addition to the well
documented importance of the disease resistance traits
of E. oleifera —such as resistance to lethal bud rot—,
if new O x G hybrids could be developed to improve
production per ha, this could help reduce the pressure to
convert biodiversity-rich tropical rainforest into oil palm
plantations. Additionally, if the introgression of the ge-
netic traits of E. oleifera to E. guineensis could produce
oils of better quality (e.g., higher oleic acid content) this
could improve the quality of oil consumed in Ecuador
and respond to demand for unsaturated oleic rich palm
oil. Despite how little is known about Ecuadorian E. olei-
fera, from this review it is clear that individuals possess
interesting agronomic traits, which are of importance to
oil palm breeders for the improvement of E. guineensis.
However, very little is known about the biodiversity of
the Ecuadorian E. oleifera populations and about what
additional genetic traits of interest could be discovered.
Questions that remain include whether other populations
of E. oleifera exist in Ecuador? What other agronomic
traits of interest are stored in the few populations that are
known in Ecuador? What are the origins of the isolated
disjointed populations in Ecuador? What we do know
is that E. oleifera is one of the most fascinating palms,
in particular due to its procumbent trunk, which gives
the impression that it moves in search of the best eco-
logical conditions in the forest. Unfortunately, in spite
of the economic importance and potential of Ecuadorian
E. oleifera, very little is known about this neglected palm
species.
ACKNOWLEDGEMENTS
Photos for Figure 1 were kindly provided by Henrik
Balslev from the AAU Herbarium (Aarhus University,
Denmark). Fieldwork for studies on E. oleifera-Taisha
fruit abscission was supported with the assistance of
Roberto Poveda, Renato Sánchez, Ramiro Zambrano
Figure 2 Examples of cultivated E. oleifera-Taisha material planted by Danec S.A. and partners in Quinindé, Ecuador. (A) An adult
individual; (B) Male inorescence at later stage of development; (C) Female inorescence; (D) Fruit bunches at dierent ripening
stages; (E) E. oleifera-Taisha individual TA26-11 does not abscise its fruit and eventually seedlings emerge from fruit that are still
attached to the fruit bunch; (F) Fruit spikelets and fruit used for phenotype test of tree TA26-11; (G) Phenotypic test with fruit bases
from tree TA26-11 before and (H) after test that shows that only one out of 15 fruit bases partially separated in the abscission zone
after 24 hours. Phenotype test was performed as described previously (Fooyontphanich et al. 2016).
17
REVISTA ECUATORIANA DE MEDICINA Y CIENCIAS BIOLOGICAS
Elaeis oleifera: a neglected palm from Ecuador
Montúfar et al.
and Angel Rubén Moreira at PDA/Murrin Companies
(Danec S.A. group), Colé (Quinindé, Ecuador). This
work was also supported by the Laboratoire Mixte
International Biodiversité des phytosystèmes Naturels et
Cultivés Andins (BIO_INCA).
REFERENCES
AAU Herbarium Database. 1990. Identication: Elaeis olei-
fera. Pedersen H.B. Specimen #97613. Aarhus University,
Denmark. Available from: http://www.aubot.dk/show_entry.
php?CatalogNumber=H.B.Pedersen97613&&sp_set=all&i-
dentication=elaeis%20oleifera
AAU Herbarium Database. 2011. Identication: Elaeis
oleifera. Balslev H., Specimen #8512. Aarhus University,
Denmark. Available from: http://www.aubot.dk/search_
results.php?collector=&number=&number_min=&number_
max=&sp_set=all&country=&family=&identification=El
aeis+oleifera&typeOf=&order=collectorReverse&order_
dir=ASC&search_log=true&Submit=search
Acosta Solís M. 1971. Palmas Ecuatorianas del Noroccidente
Ecuatoriano. Naturaleza Ecuatoriana Vol I (2): 80-163.
Arias D, Gonzalez M, Prada F, Ayala-Diaz I, Montoya C, Daza
E, Romero HM. 2015 Genetic and phenotypic diversity of na-
tural American oil palm (Elaeis oleifera (HBK) Cortés) acces-
sions. Tree Genetics & Genomes 11: 122.
Auray T, Frerot B, Poveda R, Louise C, Beaudoin-Ollivier L.
2017. Diel Patterns of Activity for Insect Pollinators of Two Oil
Palm Species (Arecales : Arecaceae). Journal of Insect Science
17(2): 45; 1-6.
Baker WJ, Couvreur TLP. 2013. Global biogeography and di-
versication of palms sheds light on the evolution of tropical
lineages. I. Historical biogeography. Journal of Biogeography
40: 274-285.
Balée W. 1989. The Culture of Amazonian Forest. Advances in
Economic Botany 7: 1-21.
Balslev H, Henderson A. 1986. Elaeis oleifera (Palmae) en-
contrada en el Ecuador. Publicaciones Museo Ecuatoriano de
Ciencias Naturales 5: 45-49.
Balslev H. 1987. Palmas nativas de la Amazonía ecuatoriana.
Colibrí 3: 64-73.
Barba J, Baquero Y, Mendoza L. 2014. Genetic Diversity of oil
palm: a source for ecological intensication of oil palm in area
aected by but rot disease. 4th International Conference on Oil
Palm and Environment (ICOPE). Available from: http://www.
palmardelrio.com/sitio/files/Presentacin_Ecupalma_Abri-
l_2014_J_Barba_PDR.pdf
Barba J, Baquero Y. 2013. Híbridos O x G obtenidos a partir
de oleíferas Taisha Palmar del Río (PDR), Ecuador. Variedad-
PDR (Taisha x Avros). Palmas 34: 315-325.
Barcelos E, Rios SD, Cunha RNV, Lopes R, Motoike SY,
Babiychuk E, Skirycz A, Kushnir S. 2015. Oil palm natural
diversity and the potential for yield improvement. Frontiers in
Plant Science 6: 190. doi: 10.3389/fpls.2015.00190
Borchsenius F, Borgtoft Pedersen H, Balslev H. 1998. Manual
to the Palms of Ecuador. AAU Report 37. Department of Sys-
tematic Botany, University of Aarhus.
Borgtoft Pedersen H, Balslev H. 1993. Palmas útiles: especies
ecuatorianas para agroforestería y extractivismo. Quito: AB-
YA-YALA. 158 p.
Bourgis F, Kilaru A, Cao X, Ngando-Ebongue GF, Drira N,
Ohlrogge JB, Arondel V. 2011. Comparative transcriptome and
metabolite analysis of oil palm and date palm mesocarp that
dier dramatically in carbon partitioning. Proceedings of the
National Academy of Sciences of the United States of America
108: 12527-12532.
Corley RHV, Tinker PB. 2016. The Oil Palm, Fifth edition.
UK: John Wiley & Sons, Ltd. The Atrium, South Gate, West
Sussex. 687 p.
Dranseld J, Uhl N, Asmussen C, Baker WJ, Harley M, Lewis
C. 2008. Genera Palmarum. The evolution and Classication
of palms. UK: Kew Publishing, Royal Botanical Gardens. 732
p.
Dussert S, Guerin C, Andersson M, Joet T, Tranbarger TJ, Pizot
M, Sarah G, Omore A, Durand-Gasselin T, Morcillo F. 2013.
Comparative Transcriptome Analysis of Three Oil Palm Fruit
and Seed Tissues That Dier in Oil Content and Fatty Acid
Composition. Plant Physiology 162: 1337-1358.
Ecocrop. 2007. Elaeis oleifera. Data Sheet. Organización de las
Naciones Unidas para la Alimentación y la Agricultura (FAO).
http://ecocrop.fao.org/ecocrop/srv/en/dataSheet?id=5632
FAOSTAT. 2018. Available from: http://www.fao.org/faostat
Fooyontphanich K, Morcillo F, Amblard P, Collin M, Jantasuri-
yarat C, Tangphatsornruang S, Verdeil JL, Tranbarger TJ. 2016.
A phenotypic test for delay of abscission and non-abscission oil
palm fruit and validation by abscission marker gene expression
analysis. Acta Horticulturae 1119: 97-104.
Guerin C, Joet T, Serret J, Lashermes P, Vaissayre V, Agbessi
MD, Beule T, Severac D, Amblard P, Tregear J, Durand-
18
REMCB 39 (1): 11-18, 2018
Gasselin T, Morcillo F, Dussert S. 2016. Gene coexpression network
analysis of oil biosynthesis in an interspecic backcross of oil palm.
The Plant Journal 87: 423-441.
Henderson A. 1995. The Palms of the Amazon. New York: Oxford
University Press. 337 p.
Henderson A, Galeano G, Bernal R. 1995. Field Guide to the Palms of
the Americas. Princeton University Press, New Jersey. 352 p.
INABIO. 2017. Agenda Nacional de Investigación sobre la
Biodiversidad. Quito: MAE, SENESCYT e INABIO. 20 p.
IndexMundi. 2018. Available from: https://www.indexmundi.com/
Lieb VM, Kerfers MR, Kronmuller A, Esquivel P, Alvarado A, Jimenez
VM, Schmarr HG, Carle R, Schweiggert RM, Steingass CB. 2017.
Characterization of Mesocarp and Kernel Lipids from Elaeis guineensis
Jacq., Elaeis oleifera [Kunth] Cortes, and Their Interspecic Hybrids.
Journal of Agricultural and Food Chemistry 65: 3617-3626.
Marin-Burgos V, Clancy JS. 2017. Understanding the expansion of
energy crops beyond the global biofuel boom: evidence from oil palm
expansion in Colombia. Energy Sustainability and Society 7: 21. DOI
10.1186/s13705-017-0123-2
Meléndez MR, Ponce WP. 2016. Pollination in the oil palms Elaeis
guineensis,. E. oleifera and their hybrids (OxG), in tropical America.
Pesquisa Agropecuária. Tropical 46 (1): 102-110.
Montúfar R, Borchsenius F, Mogollón H. 2011. Arecaceae. In: León-
Yánez S, Valencia R, Pitman N., Endara L., Ulloa Ulloa C., Navarrete
H, editors. Libro Rojo de las plantas endémicas del Ecuador, 2d. edition,
Quito: Publicaciones del Herbario QCA, Ponticia Universidad
Católica del Ecuador. p. 128-131.
Morcillo F, Gros D, Billotte N, Ngando-Ebongue G-F, Domonhédo
H, Pizot M, Cuellar T, Espeout S, Dhouib R, Bourgis F, Claverol S,
Tranbarger TJ, Nouy B, Arondel V. 2013. Improving world palm oil
production: identication and mapping of the lipase gene causing
oil deterioration. Nature Communications 4: 2160. DOI: 10.1038/
ncomms3160
Murphy D. 2006. Molecular breeding strategies for the modication of
lipid composition. In Vitro Cellular & Developmental Biology - Plant
42: 89-99.
Murphy DJ. 2009. Oil palm: future prospects for yield and quality
improvements. Lipid Technology 21: 257-260.
Murphy DJ. 2014. The Future of Oil Palm as a Major Global Crop:
Opportunities and Challenges. Journal of Oil Palm Research 26:1-24.
Ortega Cedillo D, Barrera C, Morillo E, Quintero L, Ortega JD,
Orellana J, Cevallos V, Salgado C, Souza P, Damiao C. 2016 Genetic
Diversity Within and Between Accessions of Elaeis oleifera From
the Ecuadorian Amazon. International Journal of Agriculture and
Environmental Research 2(5): 1480-1493.
Renner S. 2004. Plant dispersal across the tropical Atlantic by wind
and sea currents. International Journal of Plant Sciences 165: S23-S33.
Singh R, Low ETL, Ooi LCL, Ong-Abdullah M, Ting NC, Nagappan
J, Nookiah R, Amiruddin MD, Rosli R, Manaf MAA, Chan KL, Halim
MA, Azizi N, Lakey N, Smith SW, Budiman MA, Hogan M, Bacher B,
Van Brunt A, Wang CY, Ordway JM, Sambanthamurthi R, Martienssen
RA. 2013a. The oil palm SHELL gene controls oil yield and encodes a
homologue of SEEDSTICK. Nature 500: 340-344.
Singh R, Ong-Abdullah M, Low ETL, Manaf MAA, Rosli R, Nookiah
R, Ooi LCL, Ooi SE, Chan KL, Halim MA, Azizi N, Nagappan J,
Bacher B, Lakey N, Smith SW, He D, Hogan M, Budiman MA, Lee
EK, DeSalle R, Kudrna D, Goicoechea JL, Wing RA, Wilson RK,
Fulton RS, Ordway JM, Martienssen RA, Sambanthamurthi R. 2013b.
Oil palm genome sequence reveals divergence of interfertile species in
Old and New Worlds. Nature 500: 335-339.
Smith N. 2015. Elaeis oleifera 32. In: Palms and People in the Amazon,
Geobotany Studies,
 Springer International Publishing Switzerland.
p. 225-234.
Tranbarger TJ, Dussert S, Joet T, Argout X, Summo M, Champion A,
Cros D, Omore A, Nouy B, Morcillo F. 2011. Regulatory mechanisms
underlying oil palm fruit mesocarp maturation, ripening, and functional
specialization in lipid and carotenoid metabolism. Plant Physiology
156: 564-584.
United States Department of Agriculture, USDA. World Agricultural
Production, Circular Series WAP, 2-18, February, 2018.
Valencia R, Montúfar R. 2013. Capítulo I: Diversidad y Endemismo.
In: Valencia R, Montúfar R, Navarrete H, Balslev H, editors. Palmas
Ecuatorianas: Biología y Uso sostenible. Quito: Publicaciones del
Herbario QCA, Ponticia Universidad Católica del Ecuador. p. 3-16.
Vijay V, Pimm SL, Jenkins CN, Smith SJ. 2016. The Impacts of Oil
Palm on Recent Deforestation and Biodiversity Loss. PLoS ONE
11(7): e0159668. https://doi.org/10.1371/journal.pone.0159668
Zeven AC. 1964. On the Origin of the Oil Palm (Elaeis guineensis
Jacq.) Grana Palynologica 5: 121-123.