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A taxonomic revision of the genus Borassus L. (Arecaceae: Coryphoideae. Borasseae) is presented. Five species are recognised: B. aethiopum from Africa and Madagascar, B. akeassii from West and Central Africa, B. madagascariensis from Madagascar, B. flabellifer from South and Southeast Asia and B. heineanus from New Guinea. Pollen morphology and leaf anatomy are examined and the taxonomic history,morphology, distribution, ecology and conservation stattusare discussed.
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A Revision of Borassus L. (Arecaceae)
Author(s): R. P. Bayton
Kew Bulletin,
Vol. 62, No. 4 (2007), pp. 561-585
Published by: Springer on behalf of Royal Botanic Gardens, Kew
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KEW BULLETIN 62: 561-586 (2007) 561
A revision of
R P.
Summary. A taxonomic revision of the genus Borassus L. (Arecaceae: Coryphoideae.
Borasseae) is
presented. Five
species are recognised: B. aethiopum
from Africa and Madagascar, B. akeassii from West and Central Africa, B.
madagascariensis from Madagascar, B. flabellifer
South and Southeast Asia and B. heineanus from
New Guinea.
Pollen morphology and leaf anatomy are examined and the taxonomic history,
morphology, distribution,
ecology and conservation stattus
are discussed.
Key words. Borasseae, Coryphoideae,
anatomy, morphology, palm, pollen, taxonomy.
Borassus L. of subfamily Coryphoideae, tribe
subtribe Lataniinae (Dransfield et
al. 2005), is
one of
most widespread genera in the
Arecaceae with a
distribution encompassing tropical Africa and
Madagascar, southern and Southeast Asia and New
Guinea (Map 1). A phylogenetic study
using nuclear
and chloroplast DNA sequences (Bayton 2005),
suggests that
Borassus is monophyletic in its
circumscription, but further commentary will be
reserved for
a future
Throughout its range,
Borassus (and particularly the
Asian palmyrah, B. fiabellifer
L.) provide a huge variety
of products of great significance to local economies.
Every part of the palm is utilised, though perhaps the
most significant
product is the sweet sap (toddy,
extracted by tapping the
or inflorescences. This is
fermented into palm wine (arrack, derived from
Arabic) or, the crude sugar (jaggery,
derived from
Portuguese) is crystallised (Morton 1988). It is
not the
intention of this account to review the current and
potential uses of Borassus and several such works
already exist (e.g. Blatter 1912; Porteres 1964; Fox
1977; Kovoor 1983; Davis & Johnson 1987; Morton
1988; Burkill, 1997). However, it is noteworthy that
despite its
economic significance, the taxonomy of
Borassus has remained problematic. Species
delimitation in
Borassus has always been challenging.
Borassus palms are
massive, with
huge stems
of up to
m tall and 1
m diameter, which makes gaining access
into the crown challenging. The leaves and
inflorescences are very large and consequently only
small parts can be collected, while essential
accompanying notes are often absent. The process of
collecting, doctumenting and preserving herbarium
specimens is extremely time-consuming. Therefore
there is a dearth of herbarium material. Existing
material is often incomplete with juvenile leaves
substituted for the inaccessible mature leaves. In
addition, all Borassus species are dioecious, decreasing
further the comparability of existing specimens.
Species delimitation in Borassus has relied upon
morphological characters that reflect the narrow range
of available herbarium material, or that can only be
determined in the field.
An additional problem with
Borassus is missing type specimens. Of the seven
species recognised in the most recent taxonomic
revision of the genus (Beccari, 1924), only three have
extant type
specimens (a fourth is lectotypified
an illustration,
Moore & Dransfield 1979).
Borassus and the type species B. flabellifer
described by Rheede tot Draakestein (1678) in the
Hortus VWalabaficis
under the names Ampana (staminate
palm) and Carimpana (pistillate palm). The description
of B. flabellifer
by Linnaeus (1753) cites Rheede's
account (among others), and the accompanying
illustrations have since been selected as the lectotype
(Moore & Dransfield 1979). Borassus in
Africa was
considered by some authors to
be only a variety
of B.
flabellifer (Kirk 1867; Warburg 1895; Drude 1896,
Dammer 1901), but a second species, B. aethiopum
Mart. was described by
Martius (1838). The debate over
whether or not to
recognise Borassus aethiopum
Accepted for publication November 2006.
Centre for
Plant Diversity & Systematics, School of Biological Sciences, The University
Reading, P0 Box 221, Reading, Berkshire,
RG6 6AS, U.K. and
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K.
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resolved, with most recent authors accepting both
species (Beccari 1914, 1924; Kovoor 1983; Dransfield
1986a). A limited molecular study by Kovoor &
Hussein (1983) came to the same conclusion.
The taxonomy of the two endemic Borassus species
in Madagascar has received recent attention
(Dransfield & Beentje 1995a; Bayton et al. 2003).
Borassus madagascariensis (Jum.
& H. Perrier) Bojer ex
& H. Perrier was first
published in 1837i in a
catalogue of the native and exotic plants of
(Bojer 1837). Bojer stated that the
palm was. native to
Madagascar and could be found in the vicinity of
Majungay (now Mahajanga), and also in the Royal
Botanic Gardens of Pamplemousses (now the Sir
Seewoosagur Ramgoolam Botanical Gardens) in
Mauritius. Bojer's account does not include a
description of B. madagascariensis and his name is
therefore a nomina nuda. Jumelle and Perrier de la
Bathie (1907) accepted Bojer's account, but reduced
the species to a variety of B. flabellifer.
They later
(1913) elevated it to specific rank and it is their
accounts that inadvertently validate the epithet at
both varietal and specific ranks. A second species of
Barassus was also described byjumelle and Perrier de
la Bathie (1913). Borassus sambiranensis Jum. & H.
Perrier is restricted to the
Sambirano Region in the
northwest of the island. In the most recent taxonomic
treatment of the
Madagascar palms, Dransfield and
Beentje (1995a) concluded that there were few
morphological characters to
distinguish the endemic
Borassus species, either from each other or from B.
aethiopum in
mainland Africa.
In 1914, Beccari published the first taxonomic
revision of Borassus. In it, he described three neNw
species, and two
new varieties of B. aethiopum.
He also
transferred Borassus machadonis Ridl. into the new
genus Borassodendron Becc. (Beccari 1914). This
account was later expanded to include the other
borassoid genera and was re-published after Beccari's
death (Beccari 1924). Beccari's treatments recognised
Borassus flabellifer as restricted to
Asia and added
Borassus sundaicus Becc. from Indonesia and Borassus
Becc. from New Guinea. The latter species is
morphologically divergent from other Borassus
species, and in some ways resembles Borassodendron
machadonis (Ridl.) Becc. (Beccari 1924).
In Africa, Beccari described B. deleb Becc. from
Sudan and recognised three varieties of Borassus
aethiopum: var. senegalensis Becc. in WVest Africa,
typified by material from modern-day Mali, var.
bagamojensis Becc. in East Africa, based on material
mainland Tanzania and var. aethiopum from
Central Africa (though the type
was from
Chlevalier described a fourth variety,
var. domesticus
Chev., from
material collected in
Mali (Chevalier &
Dubois 1938). However, this name is not valid as
there was no Latin
diagnosis. In his treatment
of the
East African palmus,
Dransfield (1986a) reducect all
African taxa to a single species, with B. deleb
and the
varieties in synonymy.
However in 1996, Ake Assi &
Guinko (1996) examined a Borassus palm from
Africa that was morphologically distinctive. Further
details were provided by Ouedraogo et
al. (2002) and
Arbonnier (2002) and the consensus of opinion was
that these palms represented B. flabellifer,
unknown in
Africa. However, this taxon was later
recognised as a newv species, B. akeassii Bayton,
Ouedraogo & Guinko (Bayton et
al. 2006).
Stem & Habit
All Borassus species have a large single stem,
branching only when damage has occurred
(Ramassamv & Kannabiran 1991). The stem is
clothed in dead leaf bases, though these later abscise
cleanly leaving prominent scars. The stem of the
African and Madagascar species is
with a prominent swelling below the crown. The
anatomy and adaptive significance of this swelling
has not been investigated, but Tuley (1995) and
Henderson (2002) suggest that it is
an adaptation to
arid environments (though ventricose stems are not
exclusive to palms from arid habitats); Barot &
Gignoux (1999) proposed that it is
connected with
onset of sexual maturity. Field observation of this
character indicates that
within a population, the
swelling is usually at a similar height in each tree,
and multiple swvellings
have been observed in some
individuals (Tuley 1995). Ventricose stems are
characteristic of a number of unrelated palm genera,
many of which occur in seasonally dry habitats.
When the
ventricose stem of
Acrocomia crspa (Kunth)
C. F.
Baker ex Becc. (syn. Gastrococos crispa (Kunth)
H. E. Moore) wvas examined, the cells within were
found to
have extremely high water content (Fisher
al. 1996).
The leaves are costapalmate; the lamina is
split to one
or half of its
radius, with up to 130 induplicate
leaflets. There is a prominent triangular cleft in the
sheath below the petiole. Towards the stem, the leaf
sheath disintegrates, forming a network of tough,
woody fibres.
The petiole is robust with a flat adaxial
surface and a rounded abaxial surface; the
can be armed with spines or unarmed. There is
considerable variation in the density, shape, size and
colour of the spines, which are influenced by light
intensity and the age of the plant (Fig. 1). Adaxial
and abaxial hastulae are usually present at the
juncture of the petiole and lamina. The lamina is
broad, undulating anld stiff
towards the base.
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InduLnentum may be present on the costa and leaf
fold ridges of both surfaces, though it erodes in older
leaves. The transverse commissural veins, which link
major parallel veins of the leaf, are distinct and
their spacing varies significantly from species to
species. The leaflet apices may be acute or cuspidate,
bifid or entire.
Leaf Anatomy
The anatomy of palm leaves was examined by
Tomlinson (1961), who noted that in the Borasseae,
there was an unusually high incidence of isolateral
leaf symmetry. Most palms have leaves with an
asymmetrical (dorsiventral) anatomy i.e. a well
defined adaxial palisade mesophyll, and stomata
primarily on the abaxial surface. However, most
Borassus species lack a clearly defined palisade, and
have stomata on both surfaces, i.e. isolateral anatomy
(Tomlinson 1961). Leaf symmetry is thouLght to
correlated with light exposure (Tomlinson 1961)
and/or aridity (Barrow 1998). In the Borasseae,
genera from exposed or arid habitats (Bismarckia
Hildebr. & H. WVendl.,
Gaertn. and Medemia
Wurttenb. ex H. Wendl.) exhibit isolateral leaf
symmetry, while those from shady, humid habitats
(Borassodendron, Lodoicea Comm. ex DC. and
SatranalaJ. Dransf. & Beentje) show dorsiventral leaf
symmetry. Latania Comm. ex Juss.,
a genus of coastal
and savannas, has a somewhat intermediate leaf
anatomy (Tomlinson 1961). Borassus reflects this
overall pattern as most species of this primarily arid
zone palm have isolateral leaf symmetry, but B.
heineanus, from the rain forests of
New Guinea, has
dorsiventral leaf anatomy.
I a I.
3 cm
Fig. 1. Borassus petiole margin armature. A Borassus heineanus
Banka s.n. (K);
B. flabellifer
al. 1
160 (K); C B.
(Madagascar), Bayton
& Ranaivojaona
51 (K);
D B.
aethiopum (Madagascar), Bayton
& Ranaivojaona
53 (K);
E B.
aethiopum (juv., Kenya), Bayton
& Obunyali
14 (K);
B. aethiopum (Kenya),
4811 (K);
G B.
Faso), Bayton
et al.
72 (K);
H B.
akeassii (Burkina
Faso), Bayton
et al.
74 (K). DRAWN
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Botanic Gardens, Kew, 2007
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Borassus is pleonanthic and dioecious though
monoecy occurs rarely as a monstrosity (Louis &
Mary 1975). The inflorescences are axillary, shorter
than the leaves and the staminate and pistillate forms
are superficially dissimilar. The prophyll is the first
bract on the inflorescence and is large, bicarinate
and coriaceous, however it is rarely collected duLe to
its position deep within the leaf axils. NuLmerous
large, primary bracts sheath the inflorescence, and
these are largely indistinguishable from the
They often have hard, woody apices, fibrous margins
and green longitudinal stripes. Staminate
inflorescences are usually branched to one or two
orders (Figs 3E, 5E), but pistillate inflorescences are
typically spicate (Figs 4A, 5G).
The first-order branches of the staminate
inflorescence terminate in 1
- 3(- 5) rachillae that
are covered in scale-like rachilla bracts (Figs 3E, 5F).
These bracts are imbricate and partially connate and
consequently, the rachillae resemble massive catkins.
In the axil of each scale is a cincinnus (Fig. 3H)
containing several staminate flowers that are
obscured in a pit formed by the bracts (Figs 3F - G).
Each flower is
exserted from its pit sequentia'ly, by
elongation of the floral receptacle. Pistillate
inflorescences of most Borassus species are typically
spicate, though branching is
common in
B. akeassii
(Fig. 5H), perhaps due to human-inflicted damage
(Bayton et
al. 2006). The flower-bearing portion of
the pistillate inflorescence is covered with large
cupular bracts. The first few bracts are empty, but
subsequent bracts subtend pistillate flowers.
the staminate flowers, pistillate flowers are solitary
and superficial on the rachis.
The staminate and pistillate flowers are widely
divergent. Staminate flowers (Fig. 3J
- K) are less
than a centimetre long and are subtended by
membranous bracteoles. The calyx is angular and
membranous with three lobes. The receptacle and
corolla are fused for
most of the length of the flower,
though there are three small, keeled corolla lobes.
There are six stamens with very short triangular
filaments and medifixed, latrorse anthers. A
pistillode of varying size is usually present. The
pistillate flowers (Fig. 4A - B) are solitary and much
larger than the staminate flowers (approximately 2 x
3 cm). They are subtended by coriaceous bracteoles,
and have three distinct imbricate sepals and three
distinct imbricate petals. A staminodal ring is
present. The ovary is tricarpellate with basal septal
nectaries and the three stigmatic areas are
superficial on the apex.
Fruits and
The fruits (Fig. 4D) are very large (uLp
35 cm long)
with a coriaceoLls epicarp, wvhich varies in colour
green to
orange to black. The sepals and petals
remain attached to the rachis and persist around the
base of the developing fruit up to and incltuding
maturity. The mesocarp is pulpy and fragrant,
containing many longitudinal fibres. There is a
separate endocarp surrounding the locule of each
carpel, and this
encloses the seed forming a pyrene
(Figs 4E - H). There are between one and three
pyrenes in each fruit
and the number of pyrenes in
the fruit has an impact on the shape and size of both
the fruit
and the pyrenes (Fig. 4C). The pyrenes are
woody and are often fibrous, and there is
a great deal
of variation in shape, size and ornamentation. Some
pyrenes are bilobed and may have a deep
longitudinal furrow that can penetrate the seed
within (Fig. 4E). Other pyrenes have a prominent
external crest, the vestigial remains of the link
between the endocarps, wvhich divided duLring
development. In some specimens of B. heineanus,
there are internal flanges of the endocarp that
penetrate the seed, though these are invisible
externally (Fig. 4H). The seed almost entirely fills the
endocarp cavity; the endosperm is
homogeneouls and
bony and there is
a small central hollow. Germination
is remote-tubular, and the cotyledonary petiole
rapidly extends, penetrating the soil. The embryo is,
in effect, buried underground where it is safe from
herbivory and small fires (Tuley 1995; Barot &
Gignoux 1999).
Ferguson et
al. (1986) examined the
pollen of seven
of the eight genera of tribe Borasseae and incltuded
Borassus fiabellifer
and B. aethiopum. The eighth genus
Satranala was discovered later (Dransfield & Beentje
1995b) and its
pollen has since been examined (M.
Harley, pers. comm.). Borassoid pollen is in
monosulcate, tectate and either elliptical or
subcircular in polar view. The aperture is
almost as
long as the longest axis in
polar view (L), though in
Borassodendron it is
onlv a quarter of the length of L.
Supratectal gemmae are present in
Borassus (Fig. 2)
and Ilyphaene (Ferguson et
al. 1986). Slight differences
between the pollen of Borassus flabellifer
and B.
were noted; the gemmae of B. aethiopuum
were variable in size and a dense band of small
gemmae surrounded the aperture. In B. flabellifer
gemmae were of two
distinct sizes and the aperture
margins wvere
smooth (FerguLson
al. 1986). However,
the small sample size (one specimen per taxon)
probably precludes meaningful conclusions.
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rm S9~~
Fig. 2. Scanning
of Borassus
A B. aethiopum
Bayton & Ouedraogo 59 (K); B 8.
Brown & Brown s.n.
(K); C B. flabellifer
fox s.n. (BH);
D 8.
akeassii (Burkina
74 (K);
E B. flabellifer
Zoysa 49 (K); F B.
et al. 74 (K);
G B. heineanus
Ferrero 424 (AAU);
H B.
& Ranaivojaona
46 (K).
Scale bars
for A, B, E - H = 20 pm;
C - D = 10 pm.
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Board of Trustees
of the
Royal Botanic Gardens,
Kew, 2007
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and Ecology
A distribution map for Borassus, based on point
locality data extracted from
herbarium specimens, is
provided (Map 1). Borassus is one of the
world's most
widely distributed palm genera, challenged only
perhaps by
Calamus L., Phoenix L. or Livistona R. Br. It
occurs across the Old World tropics, but is absent
from the
Americas, except in cultivation (Morton
1988). Identifying the natural distribution of the
palm is
however difficult, due to its long history of
cultivation. Countries on the
margins of the present
distribution (e.g. Australia, Cape Verde) often have
isolated Borassus populations of unknown origin.
Moreover, some seemingly wild populations may
actually represent 'feral' crop plants. Therefore a
broad and inclusive concept of the natural
distribution of Borassus, at least in the Old World
tropics, has been accepted here.
In Africa, Borassus is restricted to the sub-Saharan
region, but is absent from large parts of the central
African rain forests, from the arid southwest and
from the Horn of
Africa. The southern limit of its
distribution is in northern South Africa. There are
several African countries where floristic data are
limited and so the presence of Borassus cannot be
determined (e.g. Angola, Burundi, Congo
Brazzaville, Liberia). Borassus has been collected on
the arid island of Soqotra, though it is probably
introduced (Miller & Morris 2004). Other African
islands where Borassus has been recorded include
Sao Tome (Exell 1944; 1973), Bioko (Guinea Lopez
1946), Cape Verde (Chevalier 1935, Henderson et
al. 2003), Pemba (Dransfield 1986a), Zanzibar
(Beentje 1990) and Mayotte (Ludwig 1999; Pascal &
Labat 2002). In
Madagascar, Borassus is restricted to
the western part of the island, and has been
recorded on the offshore islands of Nosy Be
(Bayton et al. 2003) and Nosy Mitsiou (Dransfield &
Beentje 1995a).
In Asia, Borassus flabellifer
occurs across the Indian
Subcontinent (including Sri Lanka) through
Myanmar into Indochina. Borassus occurs in China
(Yunnan), Pakistan and Peninsular Malaysia, though
is reported to have been introduced to these
countries (Whitmore 1973; Malik 1984; Pei 1991). In
Indonesia, Borassus has been recorded across the
archipelago, though excluding wetter areas such as
Borneo and the northern Moluccas. On New
Guinea, Borassus heineanus is restricted to the
northern part of the island, and is found in both
0 10002 030004000 Kilometers
Map 1. Distribution of
Borassus, based on point locality
data extracted
A B. aethiopum;
0 B. akeassii; i B.
* B. heineanus;
0 B.
Map prepared by
Justin Moat, GIS Unit, Royal Botanic Gardens,
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Papua New Gulinea and Indonesian Paptia. A few
specimens of Borassus have been found in northern
Queensland, Australia (Bailey 1902), but they are
probably not native (Jones 1984).
Borassus species occur in a variety
of habitats. Borassus
heineanus grows on alluvial sands in the tropical rain
forests of New Guinea (Barfod et al. 2001). The other
Borassus species are typical of seasonally dry open
areas with wet soils. In Africa, they are found in
mixed riverine forest where they may or may not be
the dominant species. Borassus aethiopum
can form an
almost continuous closed-canopy forest
or individuals
can be widely spaced forming distinctive palm
savannas (Tuley 1995). The accessibility of many B.
aethiopum populations and the easily-identifiable life
stages, have made the species a popular subject for
studies of population dynamics (Mordelet et
al. 1996;
Barot et al. 1999a, 1999b, 2000; Barot & Gignoux
1999; 2003). In Madagascar, Borassus exists almost
entirely in conjunction with human settlements
(Bayton et
al. 2003), though perhaps this is the result
of a shared preference for alluvial soil. In India, B.
is characteristic of sandy plains and savanna
(often near the sea), and secondary forest up to
metres altitude (Davis & Johnson 1987;
Sankaralingham & Hameed Khan 2001). On the
Indonesian island of Madura, Borassus also favours
dry sandy areas (Dransfield 1976). In many Asian
countries, Borassus is a common and familiar sight
lining the margins of rice paddies.
There are no published studies of pollination in
Borassus (Henderson 1986), though Bayton et al.
(2003) noted that
honey bees (Apis
were the
main visitors to the staminate flowers of B.
madagascariensis. Animal-mediated seed dispersal
(zoochory) in palms was reviewed by Zona and
Henderson (1989) and while there have been few
studies that included Borassus, the range of animals
implicated is impressive.
These include bats (Mtarshall
& McWilliam 1982; Marshall 1983, 1985), lions
(Dransfield 1986a), baboons (Lieberman et
al. 1979),
and African elephants (Burtt
& Salisbury 1929). While
all of these animals are known to feed on the fruit,
only in the case of elephants has effective dispersal
been demonstrated (Burtt
& Salisbury 1929). In some
areas, the
weight of the fruit
alone is the
of dispersal (barochory) (Barot et
al. 1999b).
Taxonomy & Nomenclature
In this
paper, a revision of Borassus is presented, based
on herbarium material from the following institutions:
AAU, BH, CANB, fl-G, K, NY and P. Additional
material was examined in
at the following
BM, EA, OUA and TAN. Photographs of tpe material
and other Borassius specimens from the Beccari
collection at Fl were made available by W. J. Baker and
J. Dransfield. To supplement the existing specimens,
three fieldtrips were organised: Kenya (October,
2002), Madagascar (March, 2003) and Burkina Faso
(January, 2004). Herbarium material was collected
using the modified Schweinfurth process (Dransfield,
1986b). Staminate and pistillate specimens were
collected separately with accompanying vegetati-ve
material. Access to the
was provided by use of an
extendable pole Nwith
attached pruning saw. Extensive
notes were recorded for each specimen as only small
were saved for the
herbarium. In Kenya, all of the
from three felled pistillate palms were measured
to ascertain the range of variation in fruit size. In
Burkina Faso, a sample of one hundred pistillate palms
was examined for evidence of branching in the
inflorescence. Several photographs (deposited in the
palm slide collection at K) were taken of each palm
including images of the
palm in habitat and images of
various perishable characters such as petiole and
fruit colour. Preliminary conservation status
assessments are provided folloNving IUCN guidelines
(IUCN 2001).
Ferguson et
al. (1986) examined the pollen of the
palms of tribe Borasseae and noted some differences
between Borassus flabellifer
and B. aethiopum. In an
effort to confirm these findings and identify
significant interspecific pollen characters, multiple
pollen samples were collected from all species. The
pollen was extracted from the anthers of staminate
flowers that wvere removed from herbarium
specimens, or wvere collected directly in the field.
Field-collected staminate flowers
were dried in silica
gel. The pollen was acetolysed according to the
method of Erdtman (1969). For comparative
purposes, unacetolysed pollen was also examined.
The pollen was examined using both a Reichert
Polyvar 2 light microscope and a Hitachi S-2400
scanning electron microscope. Pollen terminology
follows Punt et
al. (1994).
Leaf Anatomy
The leaf anatomy of Borassus was examined by
Tomlinson (1961), but only two
of the five
species (B.
aethiopum and B. flabellifer)
were examined. Leaf
material for anatomical study was removed from
herbarium specimens. Where possible, a single leaflet
of a similar size was used for all anatomical studies.
Square pieces of leaf approximately 1 cm2 were
sectioned (both transversely
and longitudinally) to
pm thickness using a sliding microtome. The
were stained
Alcian blue and
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and were nmounted in glycerol. The sections were
examined using a Reichert Polyvar 2 light
microscope. To clear the leaves, sections that
encompassed the whole width of one leaflet were
placed in glass dishes and covered with 5% sodium
hydroxide. The dishes were partially covered using
'cling-film' and heated in a microwave oven at full
power for five seconds. The five-second heating
periods were repeated until the leaf sections became
partially discoloured. The sections were then left in
fresh NaOH until completely discoloured. The
NaOH was then replaced with 90% bleach; when the
section was transparent, the bleach was removed and
the sections were then transferred into 30% alcohol
before being stained with safranin. The sections were
transferred into absolute alcohol via a gradient of
alcohols, and were then placed in Histoclear. The
thick midrib was removed as it prevented the
placement of a cover slip. The cleared leaflet
sections were mounted in
Canada balsam. The leaf
clearing protocol is a modified version of the
method of
Radford et
al. (1974).
Borassus L. (1753: 1183). Roxburgh (1795 - 98: 50);
Martius (1838: 219); Kunth (1841: 221); Griffith
(1851: 167); Miquel (1855: 45); Brandis (1874: 544);
Kurz (1877: 529); Bentham & Hooker (1883: 939);
Drude (1887: 40); Beccari & Hooker (1893: 481);
Baillon (1895: 255, 322); Bailey (1902: 1686); Blatter
(1912: 930); Beccari (1914: 294; 1924: 2); Backer &
Bakhuizen van den Brink (1968: 176); Malik (1984:
5); Dransfield (1986a: 18); Uhl & Dransfield (1987:
222); Pei (1991: 45); Dransfield & Beentje (1995: 51);
Dransfield & Uhl (1998: 335); Zoysa (2000: 48). Type
species: B. flabellifer
Lontarus Adans. (1763: 25). Type species: L. domestica
Massive, solitary, dioecious, pleonanthic tree palms.
Stem grey with well-defined leaf scars, base often
enlarged; stem ventricose with 1 to several swollen
sections below the crown, or not ventricose;
branching occasionally when damaged. Leaves
costapalmate, between 16 and 28 in the crown,
remaining attached to the stem in juvenile
specimens, then abscising cleanly in adulthood; leaf
sheath short and indistinct from the petiole,
splitting longitudinally to form a triangular cleft;
petiole robust, armed or unarmed, petiole colour
varying from green to yellow or black; adaxial
hastula distinct at the junction of the petiole and
lamina, rudimentary abaxial hastula present or
absent; adaxial and abaxial indumentum on some
juvenile leaves, confined to the ribs; leaflets
indtiplicate, apices bifid or entire, aculte or
cuspidate; commissuLral veins noticeable,
particularly in dried material, leaf anatomy
isolateral or dorsiventral. Staminate inflorescences
axillary, interfoliar and shorter than the leaves,
branched to
one or two orders, extended peduncle
bearing a coriaceous bicarinate prophyll;
peduncular bracts absent; rachis bracts yellow-green
with green striae, loNver rachis bracts similar to
prophyll, partial inflorescences borne in the axil of
rachis bracts, upper subtending branches
terminating in 1
- 3(- 5) rachillae; rachillae green
brown and catkin-like, covered in imbricate scale
like rachilla bracts, bracts forming pits containing a
cincinnus of 6 or more staminate flowers. Pistillate
axillary, interfoliar, pendulous, either
spicate or branched to one order; prophyll
coriaceous with a hard acuminate apex; peduncle
generally short, but sometimes extended with
several empty bracts, similar in form to the
prophyll; flower-bearing portion densely covered in
large imbricate, cutpular bracts. Staminate flowers
exserted from the pits in succession by elongation
of the floral receptacle; flowers exserted
individually or, rarely in
groups, each subtended by
membranous bracteoles; calyx membranous and
shallowly or deeply divided into three sepals,
corolla and receptacle fused for
most of the length
of the flower,
distally with three free cucullate petal
lobes; stamens 6
very short filaments, fused to
the receptacle, anthers erect, dorsifixed, latrorse;
pistillode present, small or large. Pollen
monosulcate and elliptical; longest axis in polar
view (L) 46 - 95 pm, polar axis in equatorial view
(h) 30 - 89 pim; aperture slightly shorter than the
grain (37 - 95 pim). Pistillate fowers large, solitary
and superficial on the rachis; subtended by two
large bracteoles; sepals (x 3) and petals (x 3)
similar, coriaceous and imbricate; staminodal ring
with 6 teeth usually present; ovary tricarpellate with
basal locules and orthotropous ovules, surmounted
by three stigmatic areas, septal nectaries basal.
Fruits massive and fragrant; mature fruits
may be
yellow, orange, red, green or black, produced inside
persistent perianth segments; epicarp coriaceous,
mesocarp pulpy, filled with longitudinal fibres,
endocarp woody and fibrous; pvrenes 1
- 3, pyrene
length equalling breadth or much longer than
wide, sometimes bilobed, some pyrenes have a
prominent dorsal crest, while others have
longitudinal furrows that can penetrate the seed;
additional internal flanges sometimes present in B.
heineanus. Seed closely adhering to the endocarp,
endosperm homogeneous and bony with a small
central space; embryo apical. Germination remote
tubular with a simple eophyll. n = 18 in B. flabellifer
( Read 1966;
& Devasahayam
1971) .
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There is minimal intra-specific variation in Borassus
pollen (Fig. 2) and only B. akeassii deviates significantly
from the norm. In that species, the tectum is reticulate
and the supratectal gemmae densely cover the
all but obscuring the tectum. The differences between
B. aethiopum and B. flabellifer
noted by Ferguson et al.
(1986) do not hold up to scrutiny when pollen from
larger sample of specimens is examined.
The generic name Borassus was considered
Inasculine by Linnaeus (1753) as evidenced by his use
of the masculine epithet 'flabellier'.
However, Beccari
(1914) published several new names in the genus,
including B. heinieanus,
with feminine epithets ('B.
heineana'). This error was corrected recently by
Govaerts & Dransfield (2005), but uncorrected
epithets are common in the literature.
Key to the species of Borassus
1. Stem not ventricose; petiole unarmed; leaflet apices bifid and cuspidate; lamina anatomy
dorsiventral; staminate inflorescence branched to one order; pistillate inflorescence spicate;
pistillode 0.2 - 0.5 cm long; pyrene internal flanges present or absent; New Guinea
. 1. B. heineanus
1. Stem ventricose (African species) or not (Asian species); petiole armed wvith
spines; leaflet
apices acute and entire, or splitting longitudinally with age; lamina anatomy isolateral;
staminate inflorescence branched to two
orders; pistillate inflorescence spicate or branched;
pistillode 0.01 - 0.03 cm long; pyrene internal flanges absent ................................ 2
2. Transverse commissures widely spaced (5
- 7 per cm); pistillate rachis long (? 80 cm); pollen
tectum reticulate and densely covered in gemmae; West & Central Africa . 3. B. akeassii
2. Transverse commissures closely spaced (10 - 14 per cm); pistillate rachis shorter (23 - 62 cm),
pollen tectum perforate and sparsely covered in
gemmae ................................... 3
3. Stem not ventricose; fruits black; South and Southeast Asia . 2. B. flabellifer
3. Stem ventricose; fruits green, yellow or orange; Africa & Madagascar ............................ 4
4. Petiole green along whole length, spines very snmall
(0.1 - 0.85 cm long) and pointed; fruits
green, with pointed apices; pyrenes very large (8 - 12 x 8 - 12 cm); MLadagascar * * 5. B. madagascariensis
4. Petiole black or yellow-black, spines large (0.4 - 2.8 cm long) and recurved; fruits
orange or red, with flattened or depressed apices; pyrenes large (6 - 10 x 5
- 8 cm); Africa &
Madagascar ..........................................................*4. B. aethiopum
1. Borassus heineanus Becc. (1914: 354). Beccari
(1924: 10). Type: Papua New Guinea, Sepik River,
Heine s.n. (holotype FE!).
m tall, stem diameter unknown. Leaves 20 -
28 in the crown; petiole and sheath 150 - 300 cm
long; petiole 3.1 - 5.2 cm wide at midpoint, green
with very sharp black margins, but no spines (Fig. LA);
costa 130- 150 cm long; adaxial hastula conspicuous,
to 1.2 cm, abaxial hastula absent; lamina raditus
cm maximum; leaflets 50 - 90, 3.9 - 7.1 cm wide,
apices bifid and cuspidate, shortest leaflet 110 - 130
cm long, leaf divided to
76 - 92 cm; commissural veins
3 - 6 per cm, leaf anatomy dorsiventral. Staminate
branched to one order, upper subtending
branches terminating in 1 rachilla; rachillae brown
and catkin-like, ? 70 cm long, 2.7 - 4.3 cm diameter;
rachilla bracts forming pits containing a cincinnus of
6 - 12 flowers. Pistillate inJlorescences
spicate; flower
bearing portion 37 - 49 cm long with 7 - 22 flowers
arranged spirally. Staminateflowers
exserted from pits
0.4 - 1 cm long, bracteoles 1
x 1
cm; calvx
0.3 x 0.8 cm, shallowly to deeply divided into three
sepals, petal lobes 0.2 x 0.1 cm; stamens 6 with very
short filaments, 0.11 x 0.15 cm, anthers 0.18 x 0.06
cm; pistillode distinct, 0.2 - 0.5 x 0.02 cm. Pollen
monosulcate, elliptical, 51 - 70 pm long, aperture 41 -
63 'im long, polar axis 37 - 57 pm long; tectum
perforate, sparsely covered with supratectal gemmae
(Fig. 2G). Pistillateflowers
2.5 x 2 cm; bracteoles large,
1.5 cm diam., sepals 1.5 x 2 cm, petals 1.0 x 1.5 cm.
Fruits large, 12 - 15 x 8
- 10 cm, ovoid with a slightly
pointed apex, greenish black; pyrenes 1
- 3, 9.2 - 10.5
cm x 4.8 - 7.0 cm x 4.0 - 4.5 cm; endocarp sometimes
with flanges that penetrate the seed. Figs 3
& 4.
Endemic to New Guinea. Occurring in
both Papua New Guinea and Indonesian Papua, but
restricted to the
northern side of the island.
Jayapura Regency, Bernhard Camp, Idenburg River,
April 1939, Brass 13775 (A, BRI, L); same date and
Brass 13945 (A, L) [Brass specimens verified
W. J. Baker]; Sarmi, 1
- 3 km N of Sewan on
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VOL. 62(4)
3 cm 3 cm 3 cm
30 cm 3 I
5 mm
cm ~ 1m
1 cm~~~~~~~~~~~~~5c
Fig. 3. Borassus heineanus. A costapalmate leaf; B junction of lamina and petiole
with adaxial hastula; C leaflet apex with detail
D section of unarmed petiole; E partial staminate inflorescence with detail of rachilla surface; F staminate rachilla in transverse
section; G staminate rachilla in
section; B staminate cincinnus; J staminate flower; K staminate flower in
(note elongated pistillode). A & E from photographs; B - C,
525 (AAU); D & F - K,
s.n. (K). DRAWN
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E /~~~~4c
cm V 4 cm
3 cm ~ ~ ~ ~ ~ ~ * c
Fig. 4. Borassus
heineanus. A pistillate
B pistillate
C immature
mature fruit;
E pyrene;
G pyrene
(no internal
walls); H pyrene in
A - C & G
525 (AAU);
D -F & H
420 (AAU).
Trustees of
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VOL. 62(4)
River, 1
June 1993 (9), McDonald
& Ismail
East Sepik Prov.:
Angorain sub-district, ca. 5 miles N of Timbunke on
track to
Kwoiwut, 12 Sept. 1959 (9), Pullen 1719
(CANB!); 15 km
N of Sepik River, between Timbunke
& Angoram, 19 April 2000 (9), Kjalr 525 (AAU!);
Morobe Prov.: cultivated, Papua New Guinea Forest
Research Institute, Lae, Oct. 2001 (cd), Banka s.n.
(K!, LAE); West Sepik Prov.: Beisom Non by
Creek, ca. 12 km from Bitro village, 30 Nov. 1996 (9),
Ferrero 420 (AAU!); Green River, ca. 5 km downstream
from Beimap, 30 Nov. 1996 (seedling), Ferrero
(AAU!); Green River District, Bitro village, 30 Nov.
1996 (9 & Cr), Ferrero
423, 424 (AAU!)
Tropical rain forest on alluvial sands (Barfod
et al. 2001). The palm may also be cultivated (Kjzer
CONSERVATION STATUS. Data deficient. Borassus
heineanus has been collected in only seven locations
in New Guinea. Kjaer (2003) reported a stand of
approximately 300 x 300 metres in an area by the
Sepik River in Papua New Guinea. Additional
localities may exist, and further collections are
After Georg Heine, administrator with
the German New Guinea Company, who collected
the type specimen.
VERNACULAR NAMES. Beiwof (Apau), Lipmemon
(Kamangauwi dialect).
USES. Very little is known about the uses of B.
heineanus, though Kjoer (2003) reports that the leaves
are used for thatch.
NOTES. Borassus heineanus is by far the most distinctive
species in the genus and this led Beccari (1924) to
speculate that it should be transferred into
Borassodendron. Both B. heineanus and Borassodendron
occur in humid rain forest, gene'rally an atypical
habitat for
Borassus. Unlike all other Borassus species,
the petiole is always unarmed, the leaves have a
dorsiventral leaf
anatomy and cuspidate leaflet apices,
the staminate inflorescence branches to one order
only and the staminate flowers have a large pistillode.
However, the pollen is
very similar to that found in
the other Borassus species; it has a sulcate aperture
that is
almost the length of the grain and a perforate
tectum with sparse supratectal gemmae (Fig. 2G). In
contrast, the pollen of Borassodendron has a porate
aperture and no gemmae (Ferguson et al. 1986).
Preliminary results from the molecular study confirm
the monophyly of Borassus in its current
circumscription (including B. heineanus) and suggest
Borassodendron is sister to Borassus (Bayton 2005).
The pyrenes of Borassus heineanus are rather
unusual for the genus. In most Borassus species,
pyrene length and breadth are similar or equal, but
in B. heineanus, the pyrenes are much longer than
wide (Fig. 4E). In some specimens, the endocarp has
a number of internal flanges that
penetrate the seed
(Fig. 4H). The flanges are perpendicullar to the
endocarp wall and are only visible
when the
pyrene is
viewed in transverse section. A sectioned pyrene in
the type specimen does not have internal flanges.
This character distinguishes Borassodendron (with
internal flanges) from all other Borassus species and
could indicate that a second unidentified Borassus
species occurs in New Guinea. However, no
additional morphological characters could be found
to distinguish between specimens with or without
internal flanges. The adaptive significance of these
flanges is uncertain. While living material of B.
heineanus has never been collected outside New
Guinea, two endocarps (Degener
& Degener 24625,
BH) found on a beach on the Pacific island of
Canton (Phoenix Is., Kiribati) were identified as
belonging to Borassus (Degener & Degener 1974)
and are probably attributable to this species. The
endocarps are rather long and narrow and have
perpendicular internal flanges (see Gunn & Dennis
1976: 177). These characters set them apart from all
other Borassus species. However, typical
endocarps of
B. heineanus have a small hole in the
apex to
allow the
cotyledonary stalk to exit. The Canton endocarps
have deep V-shaped clefts at the apex, usually filled
with black fibres (fibres eroded in one pyrene).
There is a great deal of natural variation in the
endocarps of better-known Borassus species. Perhaps
when more material is
collected in
New Guinea, the
presence of internal flanges or apical V-shaped clefts
in the endocarp will fit into the natural range of
variation exhibited by
B. heineanus.
2. Borassus flabellifer L. (1753: 1187). Bailey (1902:
1686); Cooke (1907: 811); Blatter (1912: 930);
Beccari (1914: 304; 1924: 4); Gamble (1922: 737);
Gagnepain & Conrard (1937: 998); Malik (1984: 5);
Pei (1991: 46); Hodel (1998: 20); Zoysa (2000: 49).
Type: Ampana Rheede (1678: 13-14, pl. 10) and
Carimpana Rheede, (1678: 11-12, pl. 9), (Lectotype
chosen by Moore & Dransfield, 1979: 60 [Ampana
designated lectotype if only one element is
Borassuis flabelliformis
L. (1774: 827) [orthographic
variant], see note below. Roxburgh (1795 - 1798:
50); Martius (1838: 219); Kunth (1841: 222);
Miquel (1855: 45); Kurz (1877: 529). Type: as B.
Lontarus domestica Gaertn. (1788: 21). Type: as B.
flabellifer L., see note below.
Borassus tunicatus Lour. (1790: 619). Kunth (1841:
224). Pholidocarpus tunicatus (Lour.) H. WAendl. ex
B. D. Jacks. (1894: 502), see note below. Type:
'Habitat in India ad oras regnorum Decan, and
Guzerate', see note below.
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Borassus sundaicus Becc. (1914: 321). Beccari (1924:
321); Backer & Bakhuizen van den Brink (1968:
176). Type: Indonesia, cultivated, Bogor Botanic
Gardens, no date or collector (holotype Fl!).
Stem to 20 m tall, grey with
well-defined leaf scars,
ventricose, but often enlarged at the base, branching
occasionally when damaged. Leaves petiole and
sheath 150 - 180 cm long; petiole 4- 6(- 7) cm wide
at midpoint, robust, bright yellow, margins black with
short (0.3 -1.3 cm) black erose teeth (Fig. 1B); costa
60 - 110 cm long; adaxial hastula conspicuous,
abaxial hastula rudimentary; lamina radius to 150 cm
maximum, dense adaxial and abaxial- indumentum
on the ribs of some juvenile leaves, leaflets -62, 4.2 -
9.5 cm wide, apices acute and entire or splitting
longitudinally with age, shortest leaflet 13 - 39 cm
long, leaf divided to 30 - 100 cm; commissural veins
11 - 18 per cm, leaf anatomy isolateral. Staminate
inflorescences branched to two orders, upper
subtending branches terminating in 1
- 3(- 4)
rachillae; rachillae green to
brown and catkin-like, 23
- 50 cm long and 1.8 - 2.5 cm diameter, sometimes
with a mamilliform apex, rachilla bracts forming pits
containing a cincinnus of 4 - 7 flowers. Pistillate
usually spicate (branched inflorescence
pictured in the lectotype), flower-bearing portion 12
- 85 cm long with 5 - 20 flowers arranged spirally.
Staminate flowers
exserted from
pits individually,
0.24 -
0.6 cm long, bracteoles 0.4 - 0.7 x 0.1 - 0.3 cm, calyx
0.3 x 0.15 cm and shallowly divided into three sepals,
petal lobes 0.1 x 0.1 cm; stamens 6
with very short
filaments, 0.2 x 0.03 cm, anthers, 0.05 x 0.03 cm;
pistillode minute. Pollen monosulcate, elliptical, 48 -
95 pm long, aperture 40 - 95 pm long, polar axis 30 -
89 pm long; tectum perforate, sparsely covered with
supratectal gemmae (Figs 2C & 2E) .
x 3 cm; bracteoles large, 2 cm diam., sepals 1.5 x 2
cm, petals 1 x 1.5 cm. Fruits
massive, 8.5 - 13 x 7.5 -
16.5 cm, yellowish black, ovoid and rounded or
flattened at the apex; produced inside persistent
perianth segments; epicarp coriaceous, mesocarp
pulp yellow, pyrenes 1
- 3, 6.1 - 10.8 cm x 4.4 - 8.5
cm x 3.1 - 4.6 cm, somewhat bilobed; most pyrenes
with one or two external, longitudinal fturrows;
internal flanges absent.
South and Southeast Asia. Determining
the 'natural' distribution of Borassus flabellifer is
essentially impossible as it is a widely planted crop
plant. It is largely restricted to areas with seasonal
rainfall and ranges from western India through
Indochina to the Lesser Sunda Islands of Indonesia.
Populations in
China, Malaysia and Pakistan may be
introduced (Whitmore 1973; Malik 1984; Pei 1991).
Borassus flabellifer
was noted to
occur in
Australia by Bailey (1902). A seedling
from a small poptulation of mature palms and
cultivated in the garden of Frank L. Jardine in
Somerset on the Cape York Peninsula of northern
Queensland. Jones (1984) noted that the palm was
still present in the garden of Jardine's abandoned
house, but the original population was never located
and the natural occurrence of Borassus in Australia is
doubtful. The Borassus palms on the island of Soqotra
have tentatively
been assigned to B. flabellifer.
geographically closer to Africa and the
palms were
identified as the
African B. aethiopum by Miller &
Morris (2004). The available herbarium material
does not allow for a conclusive identification (the
diagnostic fruits
and petiole spines are missing), but
a photograph presented by Miller & Morris (2004)
shows a mature plant without a ventricose stem. This
suggests that the
palms, which were introduced to the
island, are B. flabellifer.
Chittagong, 1867
( a), JSI-IfJ s.n. (K!); Chittagong Hill Tracts,
1880, Gamble 7781 (K!); Dhaka, 25 March 1868, Clarke
6673 (Fl!). INDIA.
Orissa State: Anugul, 19 March 1903
(a'), Haines 4034 (K!); Tamil Nadu State:
Tiruchchirappalli, 4 April 1974 (9), Mathew 7145
(K!); Madras [Chennai], Kristna Distr., 1 Jan. 1883
(da), Gamble 18581 (K!); Uttar Pradesh State: Banda,
May 1901 (d), Bell 320 (K!); West Bengal State:
cultivated, Calcutta [Kolkata] Botanic Gardens, 1869
(9 & c'), Wallich 8622 (K!,
K-W!); same locality,
(9 & a'), Gage s.n. (FI!); Calcutta [Kolkata], 1881
(9), King 69 (K!). INDONESIA.
East Nusa Tenggara
Prov.: West Timor, Kupang, 1973 (9 & d), Fox s.n.
(BH!); South East Sulawesi Prov.: Pulau Buton, 2004
(9), Rustiami 222 (BO, K!). MALAYSIA.
Kelantan State:
Tumpat, 5 May 1937 (9), Corner 32784 (BH!, K!).
SINGAPORE. Cultivated, Singapore Botanic Garden (9),
Furtado s.n. (BH!); same locality (ad), Futrtado
(BH!, K!). SRI
Central Prov.: cultivated, Royal
Botanic Gardens, Peradeniya, 22 July 1986 (9 & d),
Rutherford 115 (K!); North Central Prov.: 5 km from
Anuradhapura, 19 Sept. 1989 (9 & da), Zoysa
49 (K!).
THAILAND. Unknown locality, 1859 (ad), Schomburgk
s.n. (BH!, K!); Bangkok Prov.: Bangkok, 30
May 1906
(ad), Kerr 10708 (AAU!, BM!, K!); Bangkok, 20 May
1926 (da), Marcan 2097, 2098 (BM!); 39 km S of
Bangkok, 31 July 1963 (9 & d), King 5565 (BH!);
Nonthaburi Prov.: no locality, 24 May 1997 a(d),
Niyomdham 5038 (AAU!, K!); Phichit Prov.: 81.5 km
from Nakhon Sawan on Route 117, 26 April 2002
(a), Wilkin et al. 1160 (K!); Rayong Prov.: Salut,
Klaeng, 14 March 1928 (9), Kerr 14581 (BM!);
Songkhla Prov.: Road from
Songkhla - Sathing Phra,
Nov. 1990 (9Q
& da), Barfod 41256 (AAU!); Ban si
Chai Noi, 7 March 1994 (9 & d), Barfod 45286,
45287 (AAU!). YEMEN. Soqotra, unknown locality,
1880 (da), Balfour s.n. (K!); Hadiboh Plains, 1967.
(9), Smith s.n.
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574 KEW
HABITAT. It is difficult to determinie the original
habitat of B. flabellifer
as its distribution is so heavily
influenced by man. It occurs between sea level and
800 metres, though is
more abundant at low altitude
and is particularly common in coastal areas with
sandy or alluvial soils and in areas with permanent
moisture such as flood plains and river
valleys. It
commonly grown along the
margins of rice paddies
forming one of the
most distinctive landscapes of
Southeast Asia.
Least concern. Borassusflabelfer
widely distributed and is
common in cultivation.
The abundance of products extracted from it
ensure its continued survival.
ETYMOLOGY. The epithet 'flabellifer' translates as
'producing fans' and refers to the
palmate leaves.
NAMES. Kerigi (Soqotra), Miak tan kok
(Lao), Panna-maram (Tamil, Sri Lanka), Taan, T/han
or Tan (Thai). There are many other names applied
to this
widespread species (see Kovoor 1983), but the
name palmyrah (or palmyra), derived from the
Portuguese palmeira, has become the internationally
familiar vernacular name.
Almost every part of the palmyrah palm can be
used. The wood is thought to be termite resistant
used for construction (houses, canoes, fence posts
etc.). The leaves are used for thatch,
weaving and for
making containers for some foodstuffs. The leaflets
of B. flabellifer (together with those of Corypha
umbraculifera L.) are traditionally used as a writing
surface. They are marked using a hot metal stylus
with the parallel veins providing a convenient line
upon which to
write (Sankaralingham & Hameed
Khan 2001). Palmyrah provides a variety
of foods; the
undeveloped endosperm and apical bud (palm
cabbage) are consumed and the inflorescences are
tapped for the sweet sap. This can be fermented into
palm wine or the sugar can be crystallised. On some
Indonesian islands, this sugar is the primary source of
carbohydrates (Fox 1977). For a review
of the
uses of
B. flabellifer,
see Kovoor (1983) and Mortoil (1988).
NOTES. A number of nomenclatural problems were
uncovered while investigating the synonymy of B.
The account of Borassus flabelliformis
L. in
Systema Vegetabilium (Linnaeus 1774) matches
exactly that of B. flabellifer in Species Plantarumr
(Linnaeus 1753), and the epithets are similar,
suggesting that the former is an orthographic
variant of the latter. It could be argued that the
change in epithet was intentional, but this
render B. flabelliformis illegitimate, as both names
are based on the same type
material (Challis, pers.
comm.). When describing Lontarus domestica,
Gaertner (1788) cited the same material as
Linnaeus (1753) plus an un-named Banks
specimen, which could not be located after a
thorough search of the
Banks collection at BMN
(Vickery pers. comm.). Pholidocarpus tunicatus is
attributed to Wendland (1878: 235) in a number of
publications including Govaerts and Dransfield
(2005). However, Wendland did not make the
combination, but rather, states under Borassus
tunicata Lour., "vide Pholidocarpus" (Merrill 1935:
92). The name does not appear in the account of
Pholidocarpus, nor in the index; the combination is
therefore attributed to Jackson in Index Kewensis
(1894: 502). No original material could be found in
the Loureiro collection at BM to typify Borassus
tunicata (Vickery pers. comm.).
Borassus sundaicus was placed in synonymy with B.
flabellifer by Govaerts and Dransfield (2005), though
not noted as a new synonym. The diagnostic
characters cited by Beccari (shape of perianth
segments, endocarp crest size, seed shape) are largely
continuous when examined across the
Asian material.
The only qualitative characters cited are leaf scales
(present in B. flabellifer
and absent from B. sundaicus),
and the arrangement of perianth segments in the
fruit (imbricate in B. sundaicus, but not in B.
flabellifer). In most Borassus species, leaf scales and
indumentum are present on immature leaves, but
they erode as the leaves
mature. In addition, a degree
of overlap can be observed in the perianth segments
most Borassus species. Therefore these characters
are not taxonomically informative. Pollen from an
Indonesian specimen of B. Jfabellifer
(Fox s.n.) was
significantly larger than that from either the Thai or
Sri Lankan specimens. A sample of ten pollen grains
was measured from each specimen; the Indonesian
pollen was 70 - 95 pm long (mean 84.8 pm) while
pollen from Sri Lanka was 45 - 65 pm long (mean
53.6 pm) and pollen from Thailand was 51 - 72 pm
long (mean 60 pm). There were no noticeable
structural differences between the pollen samples.
Given the rather limited sampling of Indonesian
Borassus, it is difficult to determine whether this
aberration is
distinctive or
whether it fits within the
range of variation exhibited by B. flabellifer
Asia. However, after examination of images of the
type specimen of B. sundaicus, I can find no
additional morphological characters to distinguish
this taxon from B. flabellifer.
3. Borassus akeassii Bayton, Ouedraogo & Guinko
(2006: 354). Ake Assi & Guinko (1996: 777) [as
Borassus sp. (B. flabellifer?) ];
Arbonnier (2002: 174)
[as Borassus sp. aff.
Ake Assi et
al. (2006:
54). Type: Burkina Faso, Comoe Province, 1
km S of
Banfora, 20 Jan. 2004 (9), Bayton et
al. 69 (holotype
K!, isotypes
FTG!, OUA!).
?Borassus aethioppurn
var. domesticus
A. Chev. (1938: 103).
Type: Mali, Kaves Region: l'ile de Dinguira, Dubois
P') nom. inval., sine diagn. lat.
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Stem to 15 m tall, almost always ventricose, to
80 cm
diam., when stem marked by numerous irregular
scars, this is caused by tapping (Fig. 5A). Leaves
glaucous, 8 - 22 in the crown; petiole and sheath 90 -
160 cm long; petiole 3.0 - 7.4 cm wide at midpoint,
green, margins with small serrate black teeth (0.2 -
0.6 cm long), or teeth largely
absent (Figs 1G - H, 5B
C); costa 22 - 28 cm long; adaxial hastula
conspicuous, to
2.4 cm, abaxial hastula rudimentary;
lamina rather flat, radius to 160 cm
maximum, dense
indumentum on the ribs of some immature leaves;
leaflets 45 - 82, 2.8 - 7.3 cm wide, apices acute and
entire or splitting longitudinally with age, shortest
leaflet 58 - 147 cm long, leaf divided to 60 - 99 cm;
commissural veins 5 - 7 per cm, leaf anatomy
isolateral. Staminate inflorescences
branched to two
orders, upper subtending branches terminating in 1
- 3 rachillae; rachillae green-brown and catkin-like,
23 - 36 cm long, 2.3 - 2.5 cm diameter, usually with a
mamilliform apex; rachilla bracts forming pits that
contain a cincinnus of 5 - 10 staminate flowers.
Pistillate inflorescences
spicate or branched to one
order; rachis ? 80 cm long, flower-bearing portion 24
- 39 cm long with ? 23 flowers arranged spirally.
0.4 - 0.6 cm long, exserted from the
pits individually
or in groups of 3 - 5; bracteoles 0.8 x
0.5 cm; calyx 0.5 x 0.2 cm, shallowly divided into
three sepals, petal lobes 0.2 x 0.1 cm; stamens 6
very short filaments, 0.1 x
0.03 cm, anthers, 0.1 x 0.05
cm; pistillode minute. Pollen
monosulcate, elliptical,
51 - 72 pm long, aperture 47 - 61 pm long, polar axis
49 - 61 pm long; tectum reticulate, densely covered
with supratectal gemmae (Fig. 2D, 2F). Pistillate
3.5 x 3 cm, bracteoles 2 cm diam., sepals 2 x 2
cm, petals 1.5 x 1.5 cm. Fruits large, ? 15 cm long anci
+ 12 cm diameter, ovoid with a somewhat pointed
apex, fragrant and yellowish green at maturity,
produced inside persistent perianth segments;
pyrenes 1
- 3, 6.8 - 9.3 cm x 5.4 - 7.5 cm. Fig. 5.
DISTRIBUTION. Restricted to West and Central Africa.
Ake Assi and Guinko (1996) report that the
palm is
present in
Benin, Burkina Faso, C6te d'Ivoire, Mali,
Niger and Nigeria. Borassus akeassii (as Borassus sp.
has also been recorded in
Senegal and
the Central African Republic (Arbonnier 2002).
However, the range of B. akeassii
may be much wider
than suggested due to
misidentification of the palm
as B. aethiopum.
Johnson (1984) noted that Borassus
aethiopum in Guinea-Bissau was used for wine
production. He also noted that the palms have green,
orange-sized fruits and both these facts suggest B.
akeassii. This study can confirm the presence of B.
akeassii in Burkina Faso and Senegal only. In addition,
a specimen collected in southeastern Congo-Kinshasa
(Liben 2822) has been identified as B. akeassii. The
was based on the distinctive
commissUral veins (Bavton et
al. 2006); however, the
specimen included no fertile
Comoe Prov.:
Banfora Dept., 1 km
W of Siniana village, 20 Jan.
2004 (d), Bayton et
al., 68 (AAU!, K!, OUA!); 500 m
N of Lake Tengrela, 22Jan. 2004, Bayton et
al., 72 (K!,
OUA!); 3 km N of Banfora, 22Jan. 2004 (9 & C),
Bayton et al., 73, 74 (AAU!, K!, OUA!); Wolonkoto
village, 29 July 1998, Ouedraogo & Boussim 9 (OUA!);
Houet Prov.: Bobo-Dioulasso Dept., Dinderesso
village, 28 July 1998, Ouedraogo
& Boussim 8 (OUA!);
Kadiogo Prov.: Somgande, 18June 1998, Ouedraogo &
Guinko 2, 3 (OUA!); Ouagadougou city
centre, 4July
1998, Ouedraogo 4 (OUA!); Ouagadougou Forest, 5
July 1998, Oud,raogo 5 (OUA!). CONGO-KINSHASA.
Kasai-Occidental Prov.: Kamulolo, without date, Liben
2822 (K!). SENEGAL.
Senegambie, 1840, Brunner s.n.
(K!); Kaolack Region: Kaolack, 1906, Chevaliers.n. (P!);
Louga Region: Lingtuere Dept., Velingara in Ferlo, 30
Oct. 1989 (d), Lawesson 5444 (AAU!); Thies Region:
Mbour Dept., Ndianda village near Joal, 7 Dec. 1997
(9 & d), Ervik & Sccmbou 307 - 309 (AAU!).
Sudan savannas with 800 - 1100 mm annual
rainfall. In Burkina Faso, most populations are semi
managed for wine production. Seed is collected and
planted and the palms often have crops planted
beneath (usually cotton or cassava).
Data deficient. Borassus akeassii
may have a much wider distribution than is
known. However, in the areas where it is known to
occur, it is abundant and a significant crop for local
people. Sambou et
al. (2002; 1992) report that
exploitation is threatening populations of B.
aethiopurn in
Guinea and Senegal, but it is
that these refer to B. akeassii. In many cases, it is
difficult to determine whether populations of B.
akeassii are cultivated (i.e. planted) or merely wild
plants that
are exploited.
ETYMOLOGY. Borassus akeassii honours Professor
Laurent Ake Assi (Abidjan University, C6te d'lvoire)
who, together
Professor Sita Guinko (University
of Ouagadougou, Burkina Faso) first
the palm from B. aethiopum (Ake Assi & Guinko
VERNACULAR NAMES. Burkill (1997) lists a huge
number of indigenous names for
Borassus (ostensibly
B. aethiopum) in West Africa and many of these
be applicable to B. akeassii.
The French name r6nier
(derived from the
Wolof word r6n) is
widely used in
Francophone Africa for
both Borassus species.
USES. In Burkina Faso, the main use for the palms is
wine production, whereby the terminal bud is tapped
produce a stugary
solution that
allowed to ferment.
The undeveloped endosperm is
also consumed, and
the leaves are used for thatch
and weaving.
NOTES. Initial examination of B. akeassii suggests a
close relationship with B.
as suggested by
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cmScA SmG
Fig. 5. Borassus akeassii.
A habit,
with stem damaged by man; B petiole section without spines;
C petiole section with spines
(immature); D section of leaflet
with detail of commissures; E partial staminate inflorescence;
F partial staminate rachilla with detail
multiple flowers emerging from pits;
G partial pistillate inflorescence (flowers detached); H infructescence branched to one order. A & H
from photographs; B & D - E from Bayton et a!.
74 (K); C from Bayton et a!.
72 (K); F from
Bayton et a!.
73 (K); G from Bayton et a!. 69
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of the Royal
Botanic Gardens, Kew, 2007
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previotus authors (Ake Assi & GuLinko 1996;
Ouedraogo et
al. 2002). The similarities are largely a
matteir of scale, as B. akeassii is considerably smaller
than other African Borassus species. This has been
attributed to the intensive extraction of sap leading
to reduced growth (Porteres 1964). The small leaves,
petiole spines, inflorescences and fruits are
reminiscent of those of B. flabellifer.
However, B.
akeassii has a ventricose stem and that character is
restricted to African Borassus.
The rate of occurrence of branched pistillate
inflorescences is much higher in B. akeassii than in
any other species. Branching may occur in the
rachis towards the base of the inflorescence, or in
the flower-bearing rachillae. This may also be an
artefact of husbandry as wine tapping damages the
apical bud.
Further investigation has revealed a number of
characters that are unique to this species and
probably independent of husbandry. The pollen is
particularly distinctive (Fig. 2D, 2F); the gemmae
are denser and almost completely obscure the
tectum whereas it is clearly visible in the pollen of
all other Borassus species. In the few areas where the
tectum can be seen, it is reticulate rather than
perforate. The commissural veins in the leaves are
spaced further apart than in any species except B.
heineanus. This character can be used to identify
sterile herbarium material of B. akeassii. As the leaf
desiccates, the lamina collapses onto the network of
veins; in the leaves of B. flabellifer
and B. aethiopum
the commissures are close together and suipport the
lamina, resulting in a smooth surface. In B. akeassii,
the commissures are widely-spaced so the lamina
sinks into the rectangular pits formed bv the veins
resulting in a puckered appearance (Bayton et al.
2006). The fruits are yellow-green with a pointed
apex, thus resembling those of B. madagascariensis,
but they
are considerably smaller. These characters
should ensure that living specimens can be
identified to
species, whether they have been tapped
or not.
It is conceivable that B. aethiopum var. domesticus
could represent B. akeassii,
as the fruits
are described
Chevalier as small, globose, and yellowish-green in
colour. His choice of epithet reflects the importance
of this
variety to local people (Chevalier & Dubois
1938), which is
mirrored by B. akeassii. However, the
of var. domesticus consists of pyrenes only, and
these cannot be identified with certainty. The
presence of a second species in
West Africa could
cause some nomenclatural difficulties as the type
B. aethiopum is lost and was originally collected in
Africa. However, modern material collected
from the type locality of B. aethiopum can be
identified as that species.
4. Borassus aethiopum Mart. (1838: 221). Kunth
(1841: 223); Beccari (1914: 325; 1924: 6); Hutchinson
& Dalziel (1936: 392); Dransfield (1986a: 19); Bekele
Tesemma et al. (1993: 112); Tuley (1995: 20); Ake
Assi & Guinko (1996: 773); Arbonnier (2002: 173
[excluding top photograph]); Coates-Palgrave (2002:
99); -Bayton et al. (2003: 216); Ake Assi et al. (2006:
53). Type: Ghana, Thonning 258 (holotype lost, see
Hepper, 1976: 154); Ghana, Weija, 24
April 1957 (Q
& Cf), Tomlinson s.n. (neotype, chosen here, BH!).
Borassusflabellifervar. aethiopum (Mart.)
Warb. (1895:
20). Drude (1896: 120); Wright (1902: 117). Type:
as B. aethiopum
Borassus sambiranensisJum. & H. Perrier (1913: 67).
Beccari (1914: 346; 1924: 8); Jumelle (1938: 7);
Jumelle & Perrier de la Bathie (1945: 13);
Dransfield & Beentje (1995a: 54); Bayton et al.
(2003: 214). synon. nov. Type: none cited;
Madagascar, Antsiranana Prov., Ambanja, 28
March 2003 (ad), Bayton & Ranaivojaona 55
(neotype, chosen here, K!, P', TAN!).
Borassus deleb
Becc. (1914: 339). Beccari (1924: 8).
Type: Sudan, unknown locality and date (9),
Salvago Raggi s.n. (holotype FI').
Stem to 25m tall, almost always ventricose, to 80cm
diameter. Leaves 18 - 27 in the crown; petiole and
sheath 130 - 220 cm long; petiole 3.2 - 9 cm wide,
with large (0.4 - 2.8 cm long), recurved black
on the
margins (Fig. 1D - 1F), yellow-orange in
the distal portion, darkening to
brown-black towards
the trunk, though in immature plants, the
petiole may
be completely black; petiole spines extending along
margin of the first
costa 80- 170 cm long;
adaxial hastula conspicuous, to 2.9 cm high, abaxial
hastula rudimentary; lamina radius to 190 cm
maximum, dense indumentum on the ribs of some
immature leaves; leaflets 86 - 120, 4.7 - 11 cm
apices acute and entire or splitting longitudinally
age, shortest leaflet
45 - 92 cm long, leaf
divided to 58
- 95 cm; commissural veins 8 - 15 per cm, leaf
anatomy isolateral. Staminate inflorescences
branched to
orders, upper subtending branches terminating in
- 3 rachillae; rachillae green-brown and catkin-like,
37- 50 cm long, 2.3 - 3.8 cm diameter, sometimes
a mamilliform apex; rachilla bracts form pits that
contain a cincinnus of 8 - 14 staminate flowers.
Pistillate inflorescences
spicate; flower-bearing
portion 36
- 160 cm long
with 10 - 28 floNvers
arranged spirally.
Staminate flowers 0.2 - 0.7 cm long, exserted
individually from the pits; bracteoles 0.8 x 0.5 cm;
calyx 0.4 x 0.2 cm and shallowly divided into three
sepals, petal lobes 0.15 x 0.1 cm; stamens 6 with very
short filaments,
0.02 x
0.04 cm, anthers 0.02 x 0.05 cm;
pistillode minute. Pollen
monosulcate, elliptical, 49 -
71 um long, aperture 37 - 71 ilm long, polar axis 32 -
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61 pm long; tectum perforate, sparsely covered with
supratectal gemmae (Fig. 2A - 2B). Pistillate
3 x
3 cm, bracteoles 2 cm diam., sepals 1.5 x 2 cm and
petals 1 x 1.5 cm. Fruits
massive, 7- 17 x
7- 11 cm,
ovoid, flattened at the apex, or with a depressed apex;
fragrant, yellow to
orange or red at maturity; produced
inside persistent perianth segments; pyrenes 1 - 3, 6.4
- 10.9 cm x 5.4 - 8.0 cm x 4.2 - 5.7 cm, somewhat
bilobed; some pyrenes have one or two external
longitudinal furrows; internal flanges absent.
Found across sub-Saharan Africa as far
south as northern South Africa. Absent from
parts of
southwest and central Africa, and the Horn of
Present on a number of offshore islands including
Sao Tome and Bioko in the Gulf of Guinea, three of
the Cape Verde Islands, and Pemba, Zanzibar and
Mayotte in the
Mozambique Channel. Present in the
Sambirano region of northwest Madagascar,
including the islands of
Nosy Be and Nosy Mitsiou,
though it
may be introduced there (Bayton et al.
Prov.: Zam Dept., Rapadama, 17 Nov. 1997 (C),
Barfod 525 (AAU!); Kompienga Prov.: Pama Dept.,
Kabonga village, 1
1 Jan. 2004 (9), Bayton &
Ouedraogo 58 (K!, OUA!); Waongo village, 12 Jan.
2004 (9 & d'), Bayton & Ouedraogo 59, 60 (K!,
OUA!); Tapoa Prov.: Arli National Park, 13 Jan. 2004
(dC), Bayton
& Ouedraogo 61 (K!,
OUA!); 14Jan. 2004
(9 & C'), Bayton & Ouedraogo 62, 63 (K!, OUA!).
Kasai-Oriental Prov.: Gandajika, no
date, Liben 2817 (K!); Sud-Kivu Prov.: Impakivu
Bukavu, Kabambare, no date (d), Christiaensen 711
(K!). C6TE
Bas-Sassandra Region: Nero-Mer, 8
km NE of
Grand-Berebi, 12 Nov. 1963 (d), Oldeman
625 (K!); Lagunes Region: between Bingerville and
Potou Lagoon, 1907, Chevalier 20073 (P!); Savanna of
Cosrou, 1962 (9 & d), Leeuwenberg 4242, 4243 (K!):
Sud-Comoe Region: Assinie, Danui, 1907, Chevalier
16312 (P!). ETHIOPIA. Benishangul-Gumaz Region: W
of Gubla village, 13 Oct. 1996, Friis 7767 (K!);
Gambela Region: 5 km S of Abobo, 22
Nov. 1995 (9),
Friis 7270 (K!). GHANA. Ashanti Region: Ejura, 1921,
Chipp 773 (K!); Greater Accra Region: Accra, 1899,
Sanders s.n. (K!). KENYA.
Coast Prov.: Lamu Distr.,
Garsen (Tana River), 26 Dec. 1975 (9 & C'),
Dransfield 4810 - 4813 (EA!, K!); Kilifi Distr., Rabai,
Oct. 2002 (9 & C'), Bayton
& Obunyali 1, 2 (AAU!,
EA!, K!); Kwale Distr., Pongwe, 15 Oct. 2002 (9 &
C), Bayton & Obunyali 14 -16 (AAU!, EA!, K!);
Mivumoni, 21 Oct. 2002 (9 & d), Bayton
& Obunyali
32, 33 (EA!, K!, NY!). MADAGASCAR. Antsiranana Prov.:
4 km S of Ambanja, 24 Oct. 1988 (C'), Schatz 2415
(K!, TAN!); Ananalava, 5 km SW of
Maromandia, 4
July 1992 (9 & C), Beentje 4708, 4709 (K!, TAM!); 16
km N of Ambanja, 26 NIarch 2003, Bayton &
Ranaivojaona 53 (K!, TA.N'!); 4 km S of Ambanja, 28
March 2003 (a
C), Bayton
& Ranaivojaona 55 (K!, P!,
TAN!); Nosy Be, Hell-Ville, 29 March 2003 (9),
& Ranaivolaona 57 (K!, TANN!).
locality, 1937, Dzubois 255 (P!). NIGER. Dosso Dept.:
Dallol Foga, 1936, Auhreville s.n. (P!). NIGERIA. Delta
State: Asaba, 1906 (9), Unwin s.n. (FI!); Niger State:
Nupe, 1859 (d'), Barter 35109 (K!); SENEGAL. Thies
Region: Sebikotane, 1935, de
4484 (P!). SOUTH
AFRICA. Limpopo Prov.: Leydsdorp Distr., no date (9),
Wicht s.n. (BM!). SUDAN.
Junqali State: Central Nuer
District, 6 Oct. 1957 (d), Basinski 12 (K!); Upper
Nile State: Taufikia, April 1904 (d), Brown
& Brown
s.n. (K!). TANZANIA. Mara Region: Ndaka Plain, 20
April 1961, Greenway 12420 (K!); Pemba South
Region: Makongwe Island, 1929, Greenway 1414, 1417
(K!); Pemba Island, 19
Oct. 1931, Burtt Davy 22551
(K!); Tanga Region: Pangani, Mkwaja Ranch, 12 Sept.
1955 (d), Tanner 2187 (BH!, K!); Mweni, 3 km S of
Tanga, 5 Jan. 1976, Dransfield 4818 (K!). ZAMBIA.
Southern Prov.: Mazabuka, Nampa Estates, 7 July
1963, Rensburg 1624 (K!). ZIMBABWE.
Masvingo Prov.:
Ndanga Distr., Nltilikwe River, 12 Dec. 1953 (d),
VVild 44830, 44909 (K!).
Riverine forest and savanna in low-lying
areas, particularly on sandy or alluvial soils. Borassus
can form dense almost monospecific forest
stands or is a component of
more diverse riverine
However, it is
as the main component of palm
savannas for which it is best known. Borassus
aethiopum is well adapted to fire and herbivory and
prospers in areas with frequent burning and
browsing. After the pyrenes are dispersed, the
embryo is buried underground by the extending
cotyledonary petiole (Barot & Gignoux 1999). The
meristem is thus protected in the early years of life
during a period known as the establishment phase.
After several years surviving as a small cluster of
leaves, the palm finally generates a stem and this
grows rapidly. At this stage, the meristem is
vulnerable to fire and herbivory, but is
protected to
some extent by the skirt of dead leaves that clothes
the stem.
The dead leaf blades are highly-flammable,
but the
woody petioles are more persistent and this
may allow brush fires to
pass quickly; the leaf blade is
quickly consumed, but the petiole burns slowly,
keeping the fire
from the meristem. As a result,
the stem is covered with petioles, which may also
serve as a deterrent to large herbivores as the
are armed with spines. The accumulated leaf
sheathes and petioles are shed cleanly at maturity
(Barot & Gignoux 1999). At this
point, the
protected from fire and herbivory by
virtue of its
height above the
CONSERVATION STATUS. Least concern. Borassus
aethiopum is
widespread and common in a number of
African countries. While several populations are in
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decline (Sambou et al. 1992; 2-002), the
palm remains
common. Borassus sambiranensis is listed by the IUCN
as endangered (EN Al c) due to habitat
loss/degradation and suppression of regeneration by
fire. It is perhaps unfortunate that by placing this
taxon in synonymy
with the widespread B. aethiopunm,
the conservation status of the Madagascar
populations will be masked.
ETYMOLOGY. 'Aethiopia' was a general term used by
the ancient Greeks and Romans for
Africa south of
Libya and Egypt.
NAME. Dimaka (Sakalava, Madagascar),
Kuhe (Ashanti, Ghana), Muhama (Kibuyu, Congo
Kinshasa), Mvumo (Kiswahili, Kenya & Tanzania). A
of West African vernacular names for B. aethiopum
is provided by Burkill (1997). European names
include African fan-palm, borassus-palm, rhun-palm,
ronier (French) and palma-do-vinha (Portuguese).
USES. In some parts of Africa, Borassus populations are
'managed' by local people, while in other areas the
palm is of minimal significance. In many areas,
Borassus aethiopum is restricted to game reserves and
national parks where it is outside the reach of local
people. The leaves are used for thatch and weaving,
though species of
Hyphaene are generally preferred.
The palm is tapped for wine, though the process is
different from that
used in Asia as the apical bud is
tapped rather than the inflorescences. The fruits,
undeveloped endosperm and cotyledonary stalks are
consumed. Burkill (1997) has reviewed the uses of
Borassus aethiopum in
West Africa.
NOTES. Historically, the key question in Borassus
taxonomy has been whether to recognise both the
African B. aethiopum and the Asian B. flabellifer.
recent accounts do recognise B. aethiopum (Beccari
1924; Dransfield 1986a; 1988; Tuley 1995). The
molecular study of Kovoor & Hussein (1983) also
noted differences between the two species, thouLgh
the population sampling was extremely limited.
Qualitative characters that separate the two include
the stem (ventricose in B. aethiopum and not in B.
petiole spines (large in B. aethiopum,
B. flabellifer),
and fruit
colour (black for
B. flabellifer
and yellow-orange for
B. aethiopum). Beccari (1924)
separated the African and Asian Borassus species
based on the
degree of division of the calyx lobes of
the staminate flowers. The calyx of
Asian species was
divided to the
base, while the calyx of
African species
was only divided to the middle. This character
appears to be nothing more than an artefact of
preparation. The calyx of most staminate flowers (of
either Asian or African species) is divided to the
middle. However, the transparent, membranous
partitions between the calyx lobes tear easily. The
resultant tear is perfectly straight, giving the
impression that the calyx is divided to the
Dransfield (1986a) noted that B. aethiopum was
generally "more massive" than B. flabellifer
and for
several characters, this does appear to be the case.
The leaves have wvider
petioles with longer spines and
more leaflets, and the staminate and pistillate
rachillae are longer with more flowers. However,
while B. aethiopum and B. flabellifer
are at opposite
ends of the size range for some characters, other
Borassus species are intermediate and no truly
determinate quantitative characters were identified.
The Madagascar endemic B. sambiranensis
is placed
here in synonymy
with B. aethiopum as the two are
almost indistinguishable, both in the field and the
herbarium. Jumelle and Perrier de la
Bathie (1913)
describe in detail the differences between the two
Madagascar taxa and state that B. sambiranensis is
most similar to
B. aethiopum.
Beccari (1912 - 14, 1914,
1924) recognised B. sambiranensis, though only the
were available for him to
examine. He wrote
that they
resemble B. aethiopum var. bagamojensis
and differ only in that there is a deep hollow at the
apex. The morphology of Borassus pyrenes is
extremely variable and this character alone is
insufficient to
warrant continued recognition of B.
No type specimen was cited byJumelle
and Perrier de la
Bathie, though it
could be Perrier
12069 (lower Sambirano valley) (Dransfield &
Beentje 1995a). However, this
specimen could not be
located and a modern neotype, collected in the
Sambirano valley,
has been designated.
Borassus deleb was placed in synonymy
with B.
aethiopum by
Dransfield (1986a), as the diagnostic
fruit characters of the former fit
within the range of
variation exhibited by the latter. Due to their large
size and fleshy
consistency, few whole Borassus fruits
are preserved in
European herbaria. New taxa were
erected on the basis of differences between these
fruits without taking into account the full range of
natural variation. In an attempt to counter this
problem, every fruit from three
Kenyan specimens of
B. aethiopum (159 fruits total) was measured in the
The variation exhibited by those fruits already
encompassed almost all the variation in fruit size for
the whole genus. The fruit and leaf that are
illustrated as part of the type of B. deleb are not
significantly different from those of B. aethiopum.
Based on my own field observations and examination
of the type specimen, I agree with the conclusions of
Dransfield (1986a) to place B. deleb in synonymy
under B. aethiYtnsm.
A neotype has been designated for
B. aethiopum
holotype is
missing (Hepper 1976). The holotype
was collected in Ghana near Accra (Thonning &
Schumacher 1829). The neotype was also collected
near Accra and includes both staminate and pistillate
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VOL. 62(4)
5. Borassus madagascariensis (Jum. & H. Perrier)
Bojer exJu
& H. Perrier (1913: 61). Beccari (1914:
349; 1924: 9);Jumelle (1938: 7);Jumelle & Perrier de
la Bathie (1945: 12); Dransfield & Beentje (1995a:
52); Bayton et al. (2003: 212). Type: none cited;
Madagascar, Mahajanga Prov.: Marovoay, 14
2003 (9), Bayton &Ranaivojaona44 (neotype, chosen
here, K!, P!, TAN!).
Barassus madagascariensis Bojer (1837: 308) nom. nud.
Borassus flabellifer
var. madagascariensis Jum. & H.
Perrier (1907: 389). Type: as B. madagascariensis
& H. Perrier) Bojer exJum. & H. Perrier.
Stem to 25 m tall,
almost always ventricose, to 80 cm
diameter. Leaves 16 - 23 in the crown; petiole and
sheath 170 - 237 cm long; petiole 7.8 - 9.2 cm
wide at
midpoint, robust, yellow-green along the whole length,
margins with small serrate black teeth, 0.1 - 0.85 cm
long (Fig. 1C); petiole spines extending along the
margin of the
first leaflet; costa 77 - 82 cm long; adaxial
hastula conspicuous, to 1.5 cm, abaxial hastula
rudimentary; lamina radius to 100 cm maximum,
dense indumentum on the ribs of some immature
leaves; leaflets
102 - 132, 4.2 - 7.9 cm wide, apices acute
and entire or splitting longitudinally
with age, shortest
leaflet 68 - 83 cm long; leaf divided to 61 - 83 cm;
commissural veins 10
- 14 per cm, leaf anatomy
isolateral. Staminate inflorescences
branched to two
orders, upper subtending branches terminating in 1
- 3
rachillae; rachillae green-brown and catkin-like, 30 - 44
cm long, 2.3 - 3.3 cm diameter, usually with a
mamilliform apex; rachilla bracts form
pits that
a cincinnus of 8 - 12 staminate flowers. Pistillate
spicate; flower-bearing portion 76 - 90 cm
long with 7 - 15 flowers arranged spirally. Staminate
0.4- 0.7 cm long, exserted individually from the
pits, bracteoles 0.8 x 0.5 cm; calyx 0.5 x 0.2 cm,
divided into three sepals, petals 0.2 x 0.1 cm;
stamens 6 with very short filaments, 0.02 x 0.04 cm,
anthers, 0.14 x 0.1 cm; pistillode minute. Pollen
monosulcate, elliptical,
46- 65 pm long, aperture 38 -
52 pm long, polar axis 42 - 54 pm long; tectum
perforate, sparsely covered with supratectal gemmae
(Fig. 2H). Pistillate flowers 3.5 x 3 cm, bracteoles 2 cm
diam., sepals 2 x 2 cm and petals 1.5 x 1.5 cm. Fruits
massive, 25 - 35 x
7- 18.5 cm, ovoid with a somewhat
pointed apex; fragrant
and yellowish green at
produced inside persistent perianth segments; pyrenes
- 2(- 3), 8.7- 12.1 cm x 8.3 -12.3 cm x 6.3 - 7.5 cm,
unlobed or somewhat bilobed; some pyrenes
with one
or two
external longitudinal furrows; internal flanges
absent; an external, longitudinal crest is
Madagascar, where it is
found in the dry western part of the island.
Populations in
Mahajanga Province are all in close
There appears to
no overlap with the range of B. aethiopum in the
Sambirano region of northwest
Mahajanga Prov.:
Lower valley of Tsiribihina River, 1933, Humbert s.n.
(P!); Berevo sur Ranoba, Sept. 1940, Decary 15892
(P!); Manombolo River gorges, 10 km E of Bekopaka,
26 March 1995 (9), Du Puy
& Du Puy 791 (K!, TAN!);
Antamanakana, 44 km SW of
Katsepy, 24July 1995 (9
& d), Noblick 5055 (FTG!, K!, TAN!); Marovoay,
Colonial Experimental Garden, Oct. 1908, Prudhomme
s.n. (FI!); environs of
Marovoav, 16 March 2003 (0'),
Bayton & Ranaivojaona 46, 47 (K!, P!, TAN!); Lake
Amboromalandy, 18 March 2003 (9), Bayton &
Ranaivojaona 48 (K!); Ankarafantsika National Park,
Ampijeroa Forestry Station, 19 NMarch
2003, Bayton
Ranaivojaona 49 (K!, TAN!); Ankijabe village, near
Ambato-Boeny, 19 March 2003 (0d), Bayton &
50 (FTG!,
Maevatanana, 21
March 2003 (9 & d), Bayton
& Ranaivojaona 51, 52
(K!, MO!, TAN!); Toliara Prov.: Sakeny River plain,
July 1989, B. Du Puy et
al. 256 (K!, TAN!); unknown
date and locality,
Darian s.n. (BH!, K!);
Riverine forest in lowv-lving
areas, particularly
on sandy or alluvial soils. Borassus madagascariensis
does not form
monospecific forest. It is common in
several towns in the northwest of
Madagascar, where
grows in
gardens, streets and on waste ground.
CONSERVATION STATUS. Vulnerable. Dransfield &
Beentje (1995a) assessed B. madagascariensis as
vulnerable (VU Alc) due to habitat loss and the
encroachment of human settlements. Subsequent
investigation by the author can only confirm this
assessment. Of all the species of
Borassus, this
one has
the smallest range.
While the species does not appear
to be threatened by over-exploitation, as are some
populations of B. aethiopum in
mainland Africa, all of
the populations visited by the author were in close
proximity to expanding human settlements. Some
populations are actually confined to urban areas
(Bayton et
al. 2003) and the decline in
habitat quality
qualifies this
species as vulnerable.
The epithet 'madagascaniensis' translates
as 'of
VERNACULAR NAME. Dimaka (Sakalava). This name is
also used for
B. aethiopum in
While several uses have been recorded for B.
madagascariensis Uumelle & Perrier de la Bathie
1913), Bayton et
al. (2003) could find little evidence
exploitation in the northwest of the island.
NOTES. The case for continued recognition of this
species has been difficult to assess. Most of the
taxonomic accounts dealing with Borassus
madagasca7iensis focus on differentiating it from the
other Madagascar species, B. sambiranensis. Borassus
madagascariensis has close affinities
with B. aethiopum.
This is
not entirely unexpected as East Africa is the
likely origin of the seeds that colonised M Iadagascar.
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Both species are generally large in all aspects, and
both have a ventricose stem.
The differences between
the species are rather subtle and can be difficult to
interpret in the
herbarium. Borassus madagascariensis
has small petiole spines that are like saw-teeth
(Fig.1C), while the spines of B. aethiopum are large
and recurved (Fig. 1D - F). In B. madagascariensis, the
petiole remains green along its
entire length, in
high and low light environments. In B. aethiopum,
petiole is
yellow and turns
brown towards the stem. In
juvenile specimens of B. aethiopum, the petioles are
completely black (in full sun) or yellow-brown
(shade). Juvenile specimens of B. madagascariensis
have greenish-yellow petioles. The fruit apex of B.
madagascariensis is pointed and in this respect, it is
similar to that of Borassus akeassii.
The fruit apex of B.
aethiopum is flattened or depressed and an inverted
can sit
on its apex without toppling. The fruits
and seeds of B. madagascariensis are the largest in the
While the
colour characters can be difficult to
employ in the herbarium, the petiole spines are easily
identifiable. In combination with the large fruits
seeds, and a known origin in
Madagascar, this taxon
should be easily identified.
Jumelle and Perrier de la Bathie (1907, 1913) did
not cite a type specimen when describing B.
madagascariensis, so a neotype has been selected. The
neotype was collected in the town
Marovoay, at the
head of the Bay of Bombetok, which corresponds to
the locality
cited by
Bojer (1837: 308): "sur les
Marou-voai dans la baie de Bombetok".
Borassus aethiopum
var. bagamensis Becc. (1914: 337).
Beccari (1924: 7). Type: Tanzania, Tanga Region,
Bagamoyo, Zimmerman
s.n. (holotype F!) .
Borassus aethiopum
var. senegalensis
Becc. (1914: 334).
Beccari (1924: 7). Type: Mali, Kayes Region: Kayes,
Chevaliers.n. (holotype FI!).
These taxa are separated from typical Borassus
aethiopum primarily by small variations in pyrene
morphology. The type specimens include no
vegetative material. The shape and size of a pyrene is
influenced primarily by the number of pyrenes that
develop in each fruit.
Within each species there is
considerable variation in fruit
and therefore pyrene
size and overlap between species is
common. It
for this reason that
Dransfield (1986a) chose not to
recognise var. senegalensis or var. bagamojensis, but
rather to
unite them under the name B. aethiopum.
However, with the recognition of a second species in
Africa, it is
possible that
one or both of the
of B. aethiopum could instead be attributable to B.
akeassii. Therefore it seems inappropriate to
these names
Specimens of Uncertain
The following
African specimens
only and therefore cannot be confidently assigned to
African species.
BENIN. Unknown locality and date, Poisson s.n. (P!).
Unknown locality, 1923, Carolot s.n. (P!).
confluence with Pende, 5
May 1984, Fay s.n. (K!).
Northern Region, Rumphi Distr., 6
miles W
of Rumphi, 6 Nov. 1977, Pawek 13220 (K!). MALI.
Unknown locality, 1937, Dubois 252 (P!); Foret
Baoule, 1937, Dubois 401146 (P!); Sikasso Region:
Ouassoulou, 1891, Chevalier s.n. (P!). NIGERIA.
Northern Nigeria, 1964, Tuley s.n. (K!). SUDAN.
of the White Nile, 2 Feb. 1840, d'Arnaud s.n. (P!);
South Darfur State: Wadi Bullah, near Nyala, 5
1971, JLW 3136 (K!). TANZANIA.
Singida Region:
Lwumbu Valley, 21 Sept. 1927, Burtt 1659 (K!).
UGANDA. Unknown locality, 1955, Greenway s.n. (K!).
ZAMBIA. Lusaka Prov.: Chilanga Distr., Quien-Sabe
Farm, 1929, Sandwith s.n. (K!). UNKNOWN
West Africa, 1888, Tolly
158 (P!).
Borassus caudatus Lour. (1790: 619) = Arenga caudata
(Lour.) H. E. Moore (1960: 114).
Borassus dichotomus White (1839: 226) = Hyphaene
dichotoma (White) Furtado (1970: 301).
Borassus gomutus Lour. (1790: 618) = Arenga pinnata
(Wurmb.) Merr. (1917: 119).
Borassus ihur
Giseke (1792: 87) = Pholidocarpus ihur
Blume (1847:
Borassus machadonis Ridl. (1905: 203) = Borassodendron
machadonis (Ridl.) Becc. (1914: 361).
Borassus pinnatifrons
Jacq. (1797: 65) = Chamaedorea
pinnatifrons (Jacq.) Oerst. (1859: 14).
Borassus sonneratii
Giseke (1792: 88) = Lodoicea
Gmelin) Pers. ex
H. Wendl. (1878: 250).
Many thanks to Barbara Pickersgill, Bill Baker, John
Dransfield, Scott Zona and an anonymous referee for
their comments on the
manuscript. I am grateful to
the staff
and curators of
FTG, K, OUA, P and TAN for providing access to
specimens. Thank you to Lucy Smith (artwork) and
Justin Moat (map). Particular thanks to
Hind, Madeline Harley and Sue Mott. For assistance
with field
work, I thank Caleb Obunyali, Muthama
Muasya, Quentin Luke and the
National Museums of
Kenya (Kenya); Rolland Ranaivojaona, Helene
Ralimanana, Solofo Rakotoarisoa, Adam Britt and the
Parc Botanique et Zoologique de Tsimbazaza
(Madagascar); Amade Ouedraogo, Sita Guinko,
Joseph Boussim,
Ouoba and the
C The Board
of Trustees of the
Royal Botanic Gardens,
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Ouagadougou (Burkina Faso). This research was
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the Natural Environment Research Council
(NER/S/A/2001/06315). Additional funding for
work was provided by the South Florida Palm
Society (Kenya), the International Palm Society and
Central Florida Palm Society (Madagascar), the
Montgomery Botanical Center, the Arthur Hosier
Travel Award Scheme and the School of Plant Ciow
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