KEW BULLETIN VOL. 64: 67–94 (2009)
A monograph of Cyrtostachys (Arecaceae)
Charlie D. Heatubun1,4, William J. Baker2, Johanis P. Mogea3, Madeline M. Harley2,
Sri S. Tjitrosoedirdjo4 & John Dransfield2
Summary. Cyrtostachys Blume (Areceae: Arecaceae) is treated in this study as a genus of tree palms with a disjunct
distribution pattern across Malesia and consisting of seven species. Three species are newly recognised (C. bakeri
Heatubun, C. barbata Heatubun and C. excelsa Heatubun). Five previously accepted species (C. brassii Burret, C. kisu
Becc., C. microcarpa Burret, C. peekeliana Becc. and C. phanerolepis Burret) are reduced to synonymy with C. loriae
Becc. and one species (C. compsoclada Burret) is removed to Heterospathe as Heterospathe compsoclada (Burret) Heatubun, while C. ledermanniana Becc. is considered as a doubtful taxon. A determination key is presented and
detailed descriptions provided for all taxa. A phylogenetic analysis of all species in the genus was performed based
on morphological data. Despite the poorly resolved tree topologies, Cyrtostachys is resolved as monophyletic, with C.
glauca H. E. Moore as sister to all other species, and the west Malesian species C. renda Blume probably representing
a dispersal from within a Papuasian clade into the Sunda shelf. Natural history observations, including uses and
conservation status are also presented in this monograph.
Key Words. Arecaceae, Areceae, Arecoideae, Cyrtostachys, Malesia, morphology, taxonomy.
Introduction
Cyrtostachys Blume is a genus of palms described
almost 170 years ago by Blume (1838), based on the
remarkable species, C. renda Blume, the sealing wax
palm, a species well-known for its brilliant red leaf
sheaths. The genus is highly desirable from a horticultural point of view and it is also used locally for a
variety of purposes; the extraordinary disjunct distribution poses interesting biogeographic questions that
can only be addressed when its interspecific relationships
are better understood.
Cyrtostachys has a disjunct distribution pattern
(Map 1), with one species (C. renda) occurring to the
west of Wallace’s line in south Thailand, Malay
Peninsula, Sumatra and Borneo, and the remaining
six species on the island of New Guinea to the Solomon
Islands (Moore 1973; Dransfield 1981, 1987; Baker et al.
1998). Cyrtostachys belongs to subfamily Arecoideae and
tribe Areceae based on its pinnate, reduplicate leaf,
monoecy with flowers borne in triads of a central
pistillate and two lateral staminate flowers. It can be
distinguished from other genera of arecoid palm by the
strongly divaricate inflorescence branches, flowers
borne in pits along the rachillae, connate petals and
more than six stamens in staminate flowers (Uhl &
Dransfield 1987). Dransfield & Uhl (1986) placed
Cyrtostachys in its own subtribe, Cyrtostachydinae, as an
isolated genus with no clear relatives. Moreover, in most
studies (Asmussen & Chase 2001; Lewis & Doyle 2002;
Hahn 2002; Essig & Litten 2004) no clear affinities with
other palm genera have been found, apart from
establishing that Cyrtostachys belongs to the large group
of Indo-Pacific pseudomonomerous arecoid palm
genera tribe Areceae (sensu Dransfield et al. 2005,
2008). Recently, a phylogenetic analysis of all genera of
Areceae based on low-copy nuclear DNA provided
moderate support for a relationship between Cyrtostachys
and Clinostigma (Norup et al. 2006). However, further
evidence is required to substantiate this position. Thus,
in the new phylogenetic classification of the palm family
(Arecaceae), Cyrtostachys remains as an unplaced member of Areceae (Dransfield et al. 2005, 2008).
Although the genus Cyrtostachys is well circumscribed,
the species, with the exception of C. renda, are as yet
Accepted for publication July 2008.
1
Faculty of Forestry, Universitas Papua, Jl. Gunung Salju, Amban, Manokwari 98314( Papua Barat, Indonesia. e-mail: charlie_deheatboen@yahoo.com
2
Herbarium, Royal Botanic Gardens, Kew, Richmond( Surrey, TW9 3AB, UK.
3
Herbarium Bogoriense, Puslitbang Biologi LIPI, Jl. Ir. H. Juanda No. 22, Bogor( Jawa Barat, Indonesia.
4
Biology Department, Sekolah Pascasarjana Institut Pertanian Bogor, Baranangsiang, Bogor( 16680, Jawa Barat( Indonesia.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
68
KEW BULLETIN VOL. 64(1)
Map 1. Distribution map of the genus Cyrtostachys Blume.
rather poorly known. Most of these Papuasian species
were described based on single specimens (Beccari 1905,
1914b, 1923; Burret 1936, 1937, 1939; Moore 1966). No
monographic account has ever been published (Uhl &
Dransfield 1987; Barfod et al. 2001). Infraspecific taxa
have been described from cultivation (Tucker 1992;
Ellison & Ellison 2001; Waddel 2002), and the number
of these could increase in parallel to horticultural
demand. The genus is thus in need of reassessment.
In the 40 years since the last species, Cyrtostachys
glauca H. E. Moore, was published (Moore 1966) the
number of herbarium specimens of Cyrtostachys has
increased greatly, particularly due to activities related
to the Palms of New Guinea Project (Baker 2000, 2002);
the palm collecting density in the western part of New
Guinea, in particular, has increased dramatically. Now
we have sufficient materials in the herbarium, field
experience, and access to new data (anatomy and
pollen morphology) to study the genus and appreciate
the wide range of variation within it. Furthermore,
there has also been success in tracing isotypes of
Burret’s species, the holotypes of which were
destroyed during the Second World War in Berlin.
This monograph aims to provide a modern taxonomic treatment of Cyrtostachys and to compile and reevaluate published information on its systematics,
natural history, uses and conservation status.
Taxonomic History
The genus Cyrtostachys was described for the first time
by Carl Ludwig von Blume in 1838 based on C. renda
Blume (Blume 1838). Blume had intended to publish
the genus in Rumphia, but publication of his great
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
work was delayed until 1843 (Stafleu & Cowan 1976).
In notes with the protologue, he wrote: “…in Rumphia
l.c.” behind the genus name and he provided only a
brief description of the genus and species while in
Rumphia he provided a full description of the genus
and species accompanied by a plate with details of C.
renda from a specimen collected by Korthals near
Indrapura in Sumatra. The word Cyrtostachys comes
from the Greek word cyrtos (arched) and stachys (a
spike) in reference to the recurved inflorescence
(Whitmore 1973; Jones 1995).
Odoardo Beccari (1905) published the second
species of the genus, Cyrtostachys loriae Becc., based
on a Hartmann collection from the Owen Stanley
Range, Papua New Guinea, and this was followed by C.
kisu Becc. (Beccari 1914a), based on a specimen from
the Solomon Islands made by Guppy and C. peekeliana
Becc. (Beccari 1914b) from New Ireland (Papua New
Guinea) based on a collection made by Peekel. In
1923 another species was published by Beccari, C.
ledermanniana Becc. to accommodate Ledermann’s
specimen from Ettapenberg, Papua New Guinea
(Beccari 1923). Max Burret published C. brassii
Burret (Burret 1935) based on a specimen collected
by L. J. Brass from Kubuna, Central Division, Papua
New Guinea. Two more species of Cyrtostachys (C.
phanerolepis Burret and C. compsoclada Burret) were
published by Burret (1936). The last of Burret’s
species, C. microcarpa Burret, was published in 1939
and based on a L. J. Brass specimen from Lake
Daviambu, Fly River, Papua New Guinea (Burret
1939). Harold E. Moore Jr discovered and published
C. glauca H. E. Moore from Morobe district, Papua
New Guinea (Moore 1966).
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
69
Materials and Methods
Doyle 2002). Based on histological studies of the fruit
(Essig & Litten 2004) the genus has some similarities
with Hydriastele. Therefore, in our analysis we have
included Areca catechu L., Hydriastele costata F. M. Bailey,
Pinanga rumphiana (Mart.) J. Dransf. & Govaerts and
Rhopaloblaste ledermanniana Becc. as outgroups. Seventeen
morphological and palynological characters were included in the data matrix (Table 1), which was constructed in
MacClade 4.0 (Maddison & Maddison 2000). A parsimony analysis was performed using PAUP 4.0b10
(Swofford 1998). All characters were treated as unordered and were equally weighted. Support for the clades
was calculated by conducting a 10,000 replicate bootstrap analysis using the branch and bound algorithm.
Morphology
Morphology and distribution of the species were
studied based on specimens (dried and spirit-preserved
materials) deposited at A, AAU, BH, BO, BRI, FI, K,
KEP, L, LAE, MAN, PNH, SAN, and SING (Holmgren
et al. 1990). Many new specimens have been collected
recently by the authors. In particular, new specimens were
collected in western New Guinea (Indonesian province of
Papua and Irian Jaya Barat) by the first author in 1998 –
2005. Specimens were made in the field using standard
preparation guidelines proposed by Dransfield (1986).
Collections of leaf samples, inflorescence (rachillae with
flowers in bud and/or at anthesis), and mature fruits
were preserved in 70% alcohol for further studies.
Measurements were taken from spirit-preserved
material and dried herbarium specimens and from
cultivated individuals. Floral parts were measured
from spirit-preserved material or dried specimens
rehydrated by boiling.
Character List
General Habit Characters
1. Habit: solitary (0), clustering (1). This character
has been discussed in detail by Holttum (1955),
Dransfield (1978), Uhl & Dransfield (1987), Tomlinson
(1990) and Fisher & Maidman (1999) not only for
descriptive terminology, but also for developmental
physiology and its value for systematics.
2. Crown shape: spherical (0), hemispherical (1),
shuttle-cock shaped (2). The crown shape is an obvious
character and very helpful in recognising species in the
field. The crown itself is composed of many leaves and
forms a distinctive shape in outline. A spherical shape is
formed when leaves spread out in all directions,
forming a circular outline when viewed from a distance.
A hemispherical shape is formed when the lowest leaves
make angles 15° above or below the horizontal or leaf
inclination is ± 30°, taking into account the angles of
the lowest leaves either side of the horizontal. The
shuttle-cock crown arises when the inclination of the
lowest leaves is more than 30 ° above horizontal.
3. Leaflet arrangement: regular (0), irregular (1).
4. Leaflet thickness: papery (0), leathery (1).
5. Ramenta on abaxial surface of leaflet: absent
(0), present (1).
Pollen Morphology
Pollen material was obtained from dried herbarium
specimens and spirit-preserved material. Seventeen
samples were chosen to represent the wide range of
taxa in the genus Cyrtostachys. Details of pollen
preparation and methods for herbarium material
follow Harley (1990, 1996) and terminology follows
Punt et al. (1994), Harley (1999) and Harley & Baker
(2001). Microphotographs were taken using a Nikon
D100 Optiphot light microscope (LM) and a Hitachi
S-2400 scanning electron microscope (SEM).
Phylogenetic Analysis
A preliminary phylogenetic analysis of all species of
Cyrtostachys was conducted. In recent molecular phylogenetic studies, Cyrtostachys is resolved in a position sister to
a clade including Nenga and Areca (Asmussen & Chase
2001), or Areca (Hahn 2002), or Iguanura (Lewis &
Table 1. Data matrix for phylogenetic analysis of morphological and palynological data of species of Cyrtostachys and outgroups.
Inapplicable characters (–) are indicated. For polymorphic characters, both states are shown.
Characters
Terminal taxa
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
Cyrtostachys bakeri
Cyrtostachys barbata
Cyrtostachys elegans
Cyrtostachys excelsa
Cyrtostachys glauca
Cyrtostachys loriae
Cyrtostachys renda
Areca catechu
Hydriastele costata
Pinanga rumphiana
Rhopaloblaste ledermanniana
1
0
1
0
1
0
1
0
0
0
0
2
1
1
2
0
1
2
2
0
1
0
0
0
0
1
0
0
0
0
0
0
0
1
1
0
1
0
1
0
0
0
0
0
1
1
1
0
1
1
1
0
0
0
1
0
1
0
0
0
0
0
–
–
–
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
–
0
1
0
–
0
–
1
1
1
1
1
1
1
0
0
0
1
3
3
4
4
3
3
3
2
3
1
4
1
0
1
1
0
1
0=1
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
3
3
3
3
3
3
2
0
1
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
70
6. Arrangement of ramenta: separate (0), continuous/
connected (1).
7. Leaf sheath surface: not glaucous (0), glaucous (1).
8. Crownshaft colour: green (0), red (1).
General Reproductive Characters
9. Inflorescence basal branches: not strongly divaricate (0), strongly divaricate (1). The strongly divaricate
inflorescence is defined based on the angle formed
between the first primary branch and the rachis (main
axis) being more than 45°.
10. Inflorescence branching orders: spicate (0), 1
branching order (1), 2 branching orders (2), 3
branching orders (3), 4 branching orders (4).
11. Rachilla pits: superficial pits (0), deep pits (1).
12. Pistillate calyx persistent on rachilla: absent
(0), present (1).
13. Petals in male flowers: free (0), connate (1).
14. Filaments in male flowers: free (0), connate (1).
Pollen Characters
15. Pollen tectum surface: reticulate (0), perforate
(1), rugulate (2), verrucate-gemmate (3).
16. Trichotomosulcate grains: absent (0), present (1).
Seed Character
17. Seed endosperm: homogenous (0), ruminate (1).
Conservation Status
The conservation status of each species of the genus
Cyrtostachys was assessed based on the IUCN red list
categories and criteria version 3.1 (IUCN 2001).
KEW BULLETIN VOL. 64(1)
Results and Discussion
Species Delimitation
A morphological species concept (Davies & Heywood
1963; McDade 1995; Dransfield 1999) is used in this
monograph. Like Barrow (1998), we also looked at the
constancy of morphological character states both
within and between populations and recognised as
species only those smallest units which can be
diagnosed by constant character states.
Seven taxa of Cyrtostachys are recognised in this
monograph (Table 2); three species are new (C. bakeri
Heatubun, C. excelsa Heatubun and C. barbata Heatubun); five taxa (C. brassii Burret, C. kisu Becc., C.
microcarpa Burret, C. peekeliana Becc. and C. phanerolepis
Burret) are treated as new synonyms of C. loriae Becc.
and one taxon (C. compsoclada Burret) is incorrectly
described in Cyrtostachys and is transferred to Heterospathe. One species, C. ledermanniana Becc., is regarded
as an imperfectly known taxon. For full discussion see
Taxonomic Treatment.
Morphology
Habit
Of the seven species of Cyrtostachys, three species are
solitary (C. excelsa, C. loriae and C. barbata) and four
others are clustering (C. bakeri, C. elegans, C. glauca,
and C. renda). However, although generally solitary,
some populations of C. loriae in Indonesia, Province of
Papua, are clustering. Furthermore, C. renda also
displays variation in habit character, especially in
individuals in cultivation.
Table 2. Comparison of previously recognised taxa in Cyrtostachys with taxa recognised in this monograph. Synonyms names are
shown in italics.
Species of Cyrtostachys accepted in World checklist
(Govaerts & Dransfield 2005)
Species of Cyrtostachys accepted in this monograph
–
–
Cyrtostachys elegans Burret
–
Cyrtostachys glauca H. E. Moore
Cyrtostachys loriae Becc.
Cyrtostachys brassii Burret
Cyrtostachys kisu Becc.
Cyrtostachys microcarpa Burret
Cyrtostachys peekeliana Becc.
Cyrtostachys phanerolepis Burret
Cyrtostachys renda Blume
Areca erythropoda Miq.
Cyrtostachys lakka Becc.
Cyrtostachys bakeri Heatubun sp. nov.
Cyrtostachys barbata Heatubun sp. nov.
Cyrtostachys elegans Burret
Cyrtostachys excelsa Heatubun sp. nov.
Cyrtostachys glauca H. E. Moore
Cyrtostachys loriae Becc.
Cyrtostachys brassii Burret synon. nov.
Cyrtostachys kisu Becc. synon. nov.
Cyrtostachys microcarpa Burret synon. nov.
Cyrtostachys peekeliana Becc. synon. nov.
Cyrtostachys phanerolepis Burret synon. nov.
Cyrtostachys renda Blume
Areca erythropoda Miq.
Cyrtostachys lakka Becc.
Cyrtostachys lakka var. singaporensis Becc.
Cyrtostachys ledermanniana Becc.
Cyrtostachys compsoclada Burret
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
Imperfectly known (doubtful) taxon:
Cyrtostachys ledermanniana Becc.
Excluded Taxon:
Heterospathe compsoclada (Burret) Heatubun
comb. nov.
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
Species of Cyrtostachys are moderate to robust palms
with stems from 10 to 30 m high, they are entirely
unarmed, pleonanthic and monoecious palms and
grow in various conditions from tropical lowland
rainforest to montane forest (900 m a.s.l.); sometimes
they emerge above the forest canopy. The palm
architecture of this genus conforms to Corner’s model
for solitary species and Tomlinson’s model for clustering species (Hallé et al. 1976; Dransfield 1978;
Tomlinson 1990; Uhl & Dransfield 1987).
Stem
The stem of Cyrtostachys varies from moderate (5.5 –
10 cm in diameter) to robust (up to 30 or 40 cm in
diameter). Leaf scars are prominent and demarcate
conspicuous internodes that are mostly green near
stem tip (yellowish green, yellow, orange to red purple
with stripes in C. renda and its cultivars) and become
brown to greyish near the base. The stem in cross
section displays an outer cortex consisting of a thin
layer of about 4 mm and an inner cortex with vascular
bundles. As in Actinorhytis, Ptychococcus or Rhopaloblaste
(Banka & Baker 2004) the outer stem is usually very
hard while the inner part is softer with sparser fibres.
However, wood of Cyrtostachys has a lower number of
black fibres and is only used for certain construction
purposes. It cannot be used for traditional weapons
such as bows, arrow-heads, and spears (Heatubun et al.
in prep.).
Leaves
The crown consists of leaves ranging from about 6 – 7
per stem in Cyrtostachys glauca to 14 in C. loriae. There
are three types of crown outline in the genus:
spherical (C. barbata, C. glauca and C. loriae), hemispherical (C. elegans) and shuttle-cock shaped (C.
bakeri, C. excelsa and C. renda). According to Tomlinson
(1990) the shuttle-cock or feather duster crown shape
results from relatively few leaves in the crown, with the
leaf axis held at a high angle and abscission of the leaf
taking place as a unit before the axis droops. In
contrast, C. glauca has the lowest number of leaves in
the genus and has a spherical crown shape. All species
of Cyrtostachys have regularly pinnate leaves, except for
C. excelsa, in which leaflets are irregularly arranged in
1s or 2s. The inclination of leaflets, whether porrect or
pendulous, is associated with the crown outline.
Pendulous leaflets occur in spherical and hemispherical crowns and the porrect leaflets in shuttle-cock
shaped crowns, except in C. bakeri which has a shuttlecock shaped crown with pendulous leaflets (Fig. 5A),
and C. excelsa which has a shuttle-cock shaped crown
and slightly curved-porrect leaflets. The thickness and
coloration of leaflets when dried are leathery and
discolorous in general in Cyrtostachys, except in C.
glauca that has papery leaflets, and C. excelsa that has
concolorous leaflets. The petiole is almost missing to
71
short (to 10 cm long) in a few species, and elongate to
50 cm in C. renda and more than 50 cm (to 100 cm) in
C. glauca.
The long tubular leaf sheaths form a well-defined
crownshaft. The crownshaft is glaucous only in Cyrtostachys glauca. In C. renda the crownshaft is bright red to
orange, and in other species is green. The leaf sheath
is thick and sometimes has disintegrating fibres at the
margins just below the petiole.
The leaf anatomy of Cyrtostachys is discussed briefly
by Tomlinson (1961) based on his observation on C.
lakka (= C. renda), and he noted that Cyrtostachys is
readily distinguished from other Arecoid palms in
having markedly sinuous epidermal cell-walls and
having the hypodermal layer made up mostly of fibres
instead of colourless cells.
Indumentum
Indumentum in Cyrtostachys occurs on leaf sheaths,
leaves and inflorescences. A number of types of
indumentum occur on the leaves. Fine purple scales,
filamentous brown scales and brown to black thick
scales occur on leaf sheaths; thin to thick, caducous,
white, scale-like indumentum, with scattered, filamentous, brown to dark brown scales occurs on the
petiole; dense white or brown scales occur on the leaf
rachis, and inflexed, filamentous brown ramenta
occur scattered and/or continuous along the abaxial
surface of the mid-veins of the leaflets. The inflorescences carry less indumentum than the leaves,
although brown to rusty brown woolly indumentum
is present on the rachillae. The presence of beard-like
filamentous brown ramenta along the mid-vein on the
abaxial leaflet surface is a crucial character in recognising C. barbata from other species (Fig. 6C).
Inflorescence
All Cyrtostachys species have infrafoliar inflorescences
and are branched from two to four orders. The 2keeled prophyll encloses the inflorescence until leaf
fall, borne just above the winged base of the peduncle
and is tubular and lanceolate with winged margins,
splitting and soon caducous. The single peduncular
bract is similar to the prophyll and it too is soon
caducous. Inflorescences are strongly divaricate
(Fig. 1) with most basal branches rather diffuse and
spreading. The peduncle is usually very short, oval in
cross section. First order branches are robust, almost
as thick as the main axis (Barfod et al. 2001),
spreading, and with a short bare portion at the base,
then branching to produce diverging rachillae or
second-order branches; second order branches, when
not bearing flowers, also have a short bare portion and
then branch to produce rachillae. The rachillae are
elongate, cylindrical, rather robust, glabrous, papillose,
minutely roughened or indumentose, often brightly
coloured and expanding long before anthesis. The
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
72
KEW BULLETIN VOL. 64(1)
Fig. 1. Cyrtostachys loriae Becc. in Japen Island (Heatubun et al. 546) — strongly divaricate inflorescence with young fruits.
CHARLIE D. HEATUBUN.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
PHOTO:
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
rachilla bracts are low, triangular, spirally arranged,
rather crowded, each partially enclosing a shallow pit
bearing a triad of flowers.
The numbers of orders of branches of the inflorescence and the nature of pits have important taxonomic value to define species in Cyrtostachys. An
inflorescence branched to two orders is only found
in C. renda. Four orders of branching are found in C.
elegans and C. excelsa, while the other species have
inflorescences branched to three orders. There are
variations in the abundance and depth of pits from
species to species, from superficial pits in C. glauca
(Figs. 7D – E) to deep pits in C. loriae (Figs. 8D – E).
Inflorescences in Cyrtostachys change in colour as
they develop, from cream-coloured at the beginning
after the prophyll has fallen, followed by greenish to
green at flowering stage (anthesis of male flowers) to
fruiting (young fruits), then yellowish-brown to dark
brown at ripening (ripe fruits). All species follows this
sequence, except C. excelsa and C. renda. In C. excelsa,
pink to reddish-purple colour occurs after the prophyll
has fallen until the phase before anthesis of male
flowers, while C. renda has reddish-purple to blackish
rachillae at fruiting (young fruits) to ripening stage (ripe
fruits).
Flowers
In common with almost all Arecoideae, the flowers are
arranged in triads of a central pistillate flower and two
lateral staminate flowers, borne in pits along the entire
length of the rachilla (e.g. Fig. 7D). The staminate
flowers are cream-coloured to greenish-yellow in bud
and white to cream-coloured at anthesis. The staminate flowers are about the same size as or slightly
smaller than the pistillate. At the beginning of rachilla
emergence the pistillate flowers are not visible, being
obscured by the staminate buds. Once the staminate
flowers have fallen, the pistillate flowers become
visible and enlarge.
Staminate flowers have three, distinct, imbricate,
broad, strongly keeled sepals with minutely toothed
margins; the petals are about twice as long as the
sepals, united at the base to about one third their
length distally with triangular, valvate tips. Stamens
number 8 to 15, with awl-shaped filaments that are
connate basally and inflexed apically in bud, the
anthers being dorsifixed and latrorse. The connate
petals and stamens are distinguishing key characters
for separating Cyrtostachys from other genera in the
Indo-pacific pseudomonomerous Arecoid palm
clade.
Pistillate flowers are about the same size as, or
slightly larger than, the staminate flowers. Sepals
are three, distinct, rounded, imbricate, the margins
minutely toothed; petals are also three, distinct, imbricate proximally, asymmetrical, rounded with short
triangular valvate tips. There is a very low mem-
73
branous staminodal ring, bearing short truncate or
irregular triangular teeth. The gynoecium is unilocular, ellipsoidal with three short recurved stigmas
(e.g. Fig. 8H, J).
Fruits and Seeds
The fruits are small (to 16 × 6 mm), varying in shape
from ellipsoid, ovoid to obovoid with a beak to sickleshaped, green to black when ripe, and one-seeded.
The seeds are small (to 8 × 5 × 5 mm), globose to
ellipsoidal, apically attached, with an orbicular hilum,
and homogeneous endosperm with basal embryo (e.g.
Figs. 7K, 8K). There is no important taxonomic
character in the fruits and seeds useful to separate
species. The shape and size of fruits and seeds vary
greatly within individuals and populations.
Fruit histology of palms has given major insights
into palm systematics (Chapin et. al. 2001; Essig 1977,
1982, 1999; Essig & Hernandes 2002; Essig & Young
1979, 1985; Essig et al. 1999, 2001). Variations of fruit
pericarp among the representative species of Cyrtostachys have recently been studied in detail by Fred
Essig and co-workers. The genus can be characterised
by a combination of a papillate epidermis, a heavy
layer of tanniniferous/pigmented cells below the
epidermis, a system of vascular bundles with thick
fibrous sheaths with purely fibrous bundles frequently
above and below, the absence of brachysclereids, and a
very thin sclerified locular epidermis (Essig & Litten
2004). The authors found that one species, C. kisu (=
C. loriae), appears to be the most distinctive, with
relatively few, large bundles and tanniniferous tissue
throughout the pericarp.
Pollen Morphology
Pollen morphology was observed using both light
microscopy (LM) and scanning electron microscopy
(SEM) (Figs. 2 – 3). Using SEM the following details
were obtained: the tectum surface (topology) is
perforate, rugulate or in some species, verrucate
and/or gemmate. Sometimes, in LM or SEM preparations, a few still intact tetrahedral post meiotic tetrads
were observed. Pollen morphology for individual species
is included in the species descriptions.
Results from pollen morphological data provide
limited variation in characters to aid species delimitation,
especially in the Papuasian taxa. This reflects the findings
of Harley (1999), who conducted a general survey of
pollen morphology throughout the genera of the palm
family and found that pollen data are not often useful in
helping to support delimitation at the species level.
However, the data give a clear distinction between the
west Malesian Cyrtostachys renda and the Papuasian taxa,
in which the topology of the tectum surface of C. renda is
microfossulate-rugulate (Fig. 3J – M), in contrast to the
verrucate-gemmate tectum topology in the Papuasian
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74
KEW BULLETIN VOL. 64(1)
Fig. 2. Some of the pollen morphological diversity in Cyrtostachys. A – C C. bakeri (from Baker 1138): A whole grain, proximal face;
B whole grain, distal face; C group of grains to show general appearance of pollen. D, E C. barbata (from Brass 13707): D whole
grain, proximal face; E two rather collapsed grains, apparently conjoined. F – H C. elegans (from Heatubun 341): F whole grain,
proximal face; G whole grain, distal face showing trichotomosulcate aperture; H close up of verrucate surface, proximal face. J – L
C. excelsa (from Heatubun 330): J three grains adhered, showing proximal faces; K whole grain, proximal face; L close up of
verrucate surface, proximal face. Scale bar: 5 m.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
75
Fig. 3. Some of the pollen morphological diversity in Cyrtostachys (continued). A – C C. glauca (from Barfod 454): A whole grain,
proximal face; B close up of verrucate surface, proximal face; C group of grains to show general appearance of pollen. D – H C.
loriae (D, G from Barfod 463; E, F, H from Whitmore 3945): D group of grains; E whole grain, distal face showing monosulcate
aperture; F whole grain, distal face showing trichotomosulcate aperture; G two grains, showing proximal faces; H close up of
gemmate surface, proximal face. J – M C. renda (from Keith 2491): J whole grain, proximal face; K whole grain, distal face showing
monosulcate aperture with membrane semi intact; L whole grain, distal face showing trichotomosulcate aperture; M close up of
micro-fossulate surface, proximal face. Scale bar: 5 m.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
76
species (Figs. 2A – K, 3A – H). As well as Cyrtostachys,
these types of tectum surface topology are also found in
some species of other palm genera; perforate and
rugulate types are the second most widespread ectexine
type in the palms and it occurs in all subfamilies except
Nypoideae. The tectate verrucate type is uncommon
except in Calamus (Calamoideae: Calameae) where it is
noted in about seventeen species; in Coryphoideae it is
known in Hyphaene (Borasseae: Hyphaeninae) and in
Arecoideae there are three other genera where some
species have a verrucate, or regulate-verrucate pollen
surface — Chamaedorea carchensis Standl. & Steyerm.
(Chamaedoreeae), Aiphanes (Cocoseae: Bactridinae)
and Prestoea (Euterpeae). The gemmate type is one of
three ectexine types that have supratectate and intectate forms, supratectate gemmae occur on the ectexine
of five species of Calamus (Calamoideae: Calameae),
Hyphaene (Borasseae: Hyphaeninae), Borassus (Borasseae: Lataniinae), Attalea (Cocoseae: Attaleinae), Bactris
(Cocoseae: Bactridinae), Ceroxylon (Ceroxyloideae: Ceroxyleae), Dictyocaryum and Iriartea (both Arecoideae:
Iriarteeae) and Voanioala (Cocoseae: Attaleinae).
Phylogenetic Analysis
The cladistic analysis of morphological and palynological data generated eight equally most parsimonious trees (length=26 steps, consistency index=0.69,
retention index=0.79, rescaled consistency index=
0.55). A strict consensus of the trees, including
bootstrap support values, and a single tree selected
randomly from the most parsimonious trees, are shown
in Fig. 4.
The topology of the strict consensus tree is poorly
resolved and supported (Fig. 4); only three nodes
show bootstrap supports above 75%. Although the
resolution is low, the monophyly of Cyrtostachys is
well supported (bootstrap=98%) and C. glauca is sister
to all remaining species of Cyrtostachys (bootstrap=
75%). The topology of the tree suggests that C.
renda dispersed from within New Guinea/Solomon
(Papuasia) clade into the Sunda Shelf.
The low resolution of the strict consensus tree
gives clear indication of the high level of character
conflict caused by homoplasious characters. The
recovery of Cyrtostachys as a monophyletic genus
confirms again its circumscription as a distinct genus.
Several uncontradicted synapomorphies optimise at
the highly supported node (95% bootstrap support)
at the base of the genus: pistillate calyx persistent
on rachilla, petals in male flowers connate, filaments
in male flowers connate, endosperm homogeneous.
The character states “pistillate calyx persistent on
rachilla” and “endosperm homogeneous” are also
found in other genera of Arecoid palms, but not in
outgroups used in this study. Uhl & Dransfield
(1987) define Cyrtostachys as a genus separate from
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KEW BULLETIN VOL. 64(1)
other genera by the characters “connate petals
and filaments in staminate flowers,” also “strongly
divaricate inflorescence” and “flowers borne in pits
along the rachilla.” However, in this analysis these
character states are homoplasious because they are
also found in one of the outgroups (Rhopaloblaste
ledermanniana Becc.).
The support of Cyrtostachys glauca as sister to all
species of Cyrtostachys is also predicted; occasionally
there has been some doubt as to whether C. glauca is a
member of the genus Cyrtostachys because of the
nature of the pits along the rachillae. Other autapomorphic characters (leaf sheath surface glaucous and
trichotomosulcate grains absent) also contributed to
the isolation of C. glauca from other species.
Biogeography and Ecology
Cyrtostachys is a tropical genus with a disjunct distribution pattern; one species (C. renda) occurs to the west
of Wallace’s line from south Thailand to the Malay
Peninsula, Sumatra and Borneo, and the other six
species on the island of New Guinea to the Solomon
Islands. How this disjunct distribution arose has been
discussed for three decades (Moore 1973; Dransfield
1981, 1987; Baker et al. 1998).
The cladistic analysis suggests that Cyrtostachys renda
dispersed from within the New Guinea-Solomon
(Papuasia) clade into the Sunda Shelf, although there
is no evidence yet of macro fossils and/or fossil pollen
of Cyrtostachys from areas between the two distribution
regions of the genus (Van der Kaars, pers. comm.) to
support that analysis. In fact, information generally
from macro fossils and fossil pollen in the Indonesia
archipelago is rather poor, especially in the gap
between the two distribution regions of Cyrtostachys.
Four locations of excavation have been reported so
far, they are: Banda Sea (Van der Kaars et al. 2000),
Halmahera and North Moluccas Sea (Barmawidjaya
et al. 1993), and Tondano (Dam et al. 2001). Further
study is needed.
Meanwhile, New Guinea appears to be the centre
of diversity for Cyrtostachys, based on numbers of
species. Six of the seven species of Cyrtostachys
currently recognised in this monograph are originally
from New Guinea. The complex and rapid tectonic
change as well as climate and the wide range of
ecological conditions are probably the major causes of
speciation in New Guinea (Pigram & Davies 1987; Van
Welzen 1997; Bachman et al. 2004).
Cyrtostachys occurs in a wide range of ecological
conditions from swampy areas in lowlands to heath
forest in lower montane vegetation, from evergreen
rain forest to dry areas in savannah lands and from the
continental to small off-shore islands, with C. renda and
C. loriae widespread species in each region in west
Malesia and Papuasia respectively.
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
77
Fig. 4. A Strict consensus of 8 most parsimonious trees found from cladistic analysis of morphological and palynological data from
Cyrtostachys. Numbers above branches indicate bootstrap support. B One of 8 most parsimonious trees (length = 26 steps, CI =
0.69, RI = 0.79, RCI = 0.55) found from cladistic analysis of morphological and palynological data from the genus Cyrtostachys.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
78
Uses
Ethnobotanical information on Cyrtostachys, especially of the New Guinean taxa, will be discussed in
detail by Heatubun et al. (in prep.), but in general
the genus Cyrtostachys plays an important role in the
traditional life of local communities. Many parts of
the palm are utilised: the stem, leaves, leaf sheath,
cabbage (palm heart), and mid-vein of leaflets. The
species C. loriae in particular, has been widely used by
indigenous people in New Guinea through to the
Solomon Islands for construction materials in traditional house (piling, flooring, thatching), and bridge
building, as water pipes and in other local methods of
construction, such as tools, weapons and other equipment including mattresses. The remarkable species
C. renda has been established in horticulture as an
ornamental and has substantial economic importance.
Many botanic gardens and palm collectors all over the
world put this species as a priority in their collection
lists. A number of cultivars have been described from
cultivation (Tucker 1992; Ellison & Ellison 2001;
Waddel 2002).
In addition, the local names, common names,
trade names, and folklore or mythologies that we
found for the genus Cyrtostachys give proof of the
relationship between species of Cyrtostachys and the
human communities living near the habitat of this
palm.
Taxonomic Treatment
Cyrtostachys Blume (1838: 66; 1843: 101); Moore
(1973: 107); Dransfield & Uhl (1986: 8); Uhl &
Dransfield (1987: 378). Type: C. renda Blume.
Solitary or clustering, moderate to robust, unarmed,
pleonanthic, monoecious palms. Stem erect, bare,
conspicuously ringed with leaf scars, often bearing a
mass of adventitious roots at the base, where clustering, the clump rather close, or more diffusely spreading by stolons. Leaves pinnate, neatly abscising; sheath
tubular, forming a well defined crownshaft, brilliantly
orange-red coloured in one species (C. renda), glabrous or scaly; rachis like the petiole but angled
adaxially; leaflets always single-fold, acute or acuminate or sometimes rounded with a bifid tip, regularly
arranged or grouped with two leaflets in one species
(C. excelsa), often stiff, sometimes slightly paler beneath, ± glabrous adaxially, abaxially often with
ramenta along mid vein and sometimes minutely
dotted between the veins, transverse veinlets conspicuous or obscure. Inflorescence apparently protandrous, infrafoliar, branched to 2 – 4 orders, rather
diffuse and spreading; peduncle usually very short, ±
oval in cross section; prophyll enclosing the inflorescence until leaf fall, borne just above the winged base
of peduncle, tubular, 2-keeled, ± lanceolate, with
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KEW BULLETIN VOL. 64(1)
winged margins, splitting, soon caducous; peduncular
bract borne just above the prophyll, completely
enclosing the inflorescence, splitting longitudinally
like the prophyll, caducous; subsequent bract very
inconspicuous, incomplete, low, triangular; rachis
longer than the peduncle; first order branches robust,
spreading, with a short bare portion at the base, then
branching to produce diverging rachillae or secondorder branches; second-order branches, when not
rachillae, also with short bare portion and then
branching to produce rachillae; rachillae elongate,
cylindrical, rather robust, glabrous, papillate, minutely
roughened or indumentose, sometimes brightly coloured, expanding long before anthesis; rachilla bracts
low, triangular, spirally arranged, rather crowded, each
partially enclosing a shallow pit bearing a triad of
flowers, triads borne throughout the length of the
rachillae; floral bracteoles membranous, very small
and inconspicuous. Staminate flowers with 3, distinct,
imbricate, broad, strongly keeled sepals with minutely
toothed margins; petals about twice as long as sepals,
united at the base to c. ⅓ their length, distally with
triangular, valvate tips; stamens 8 – 15, the filaments
awl-shaped, connate basally, apically inflexed in bud,
anthers dorsifixed, latrorse; pollen grains ellipsoidal,
less frequently, oblate triangular, symmetric or slightly
asymmetric; aperture a distal sulcus or trichotomosulcus; ectexine tectate, perforate, rugulate, in some
species with verrucate or gemmate supratectal processes, aperture margin similar; infratectum columellate; longest axis ranging from 27 – 57 μm, shortest
axis ranging from 23 – 44 μm; post-meiotic tetrads
tetrahedral; pistillode almost as long as filament,
narrow, elongate, trifid. Pistillate flowers about same
size as or slightly larger than the staminate; sepals 3,
distinct, rounded, imbricate, the margins minutely
toothed; petals 3, slightly larger than sepals, distinct,
imbricate, proximally asymmetrical, rounded with
short triangular valvate tips; staminodal ring membranous, very low, bearing short truncate or irregularly
triangular teeth; gynoecium unilocular, ellipsoidal
with 3 short recurved stigmas, ovule pendulous from
apex of locule, form unknown. Fruits 1-seeded, broad
to narrow-ellipsoidal or sickle-shaped, usually black,
the perianth whorls persistent, stigmatic remains
apical; epicarp smooth, contrasting with rachilla,
mesocarp thin, closely adhering to the seed; pericarp
anatomy: combination of papillate epidermis, heavy
layer of tanniniferous/pigmented cells below the
epidermis, a system of vascular bundles with thick
fibrous sheaths with purely fibrous bundles frequently
above and below, absence of brachysclereids, and a
very thin sclerified locular epidermis. Seed globose or
ellipsoidal, apically attached, the hilum orbicular,
endosperm homogenous; embryo basal. Germination
adjacent-ligular; eophyll bifid with narrow lobes.
Cytology n = 16 (C. renda, Sarkar 1970).
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
79
Key to Species of Cyrtostachys
1. Crownshaft and leaf sheath orange to bright red; West Malesia (Southern Thailand, Malay Peninsula, Sumatra,
Borneo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. C. renda
Crownshaft and leaf sheath glaucous and green; Papuasia (New Guinea to the Solomon Islands . . . . . . . . . . . . . 2
2. Crownshaft and leaf sheath glaucous; elongate petiole (25 – 100 cm long); inflorescence slender, whitish when
dried, and rachilla with superficial pits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. C. glauca
Crownshaft and leaf sheath green; petiole relatively short (less than 20 cm long); inflorescence relatively large
to robust, not whitish when dried, and rachilla with deep pits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaflets irregularly arranged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. C. excelsa
Leaflets regularly arranged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Leaflets pectinate with beard-like ramenta continuous along mid-vein on abaxial surface of leaflets . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. barbata
Leaflets regularly pinnate, ramenta rare on abaxial surface of leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Stem solitary; crown spherical in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. C. loriae
Stem clustered; crown not spherical in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Crown shuttle-cock shaped in outline; inflorescence compact, branched to 3 orders; stamens 12 . . . . 1. C. bakeri
Crown hemispherical in outline; inflorescence elongate, branched to 4 orders; stamens 9 . . . . . . . . 3. C. elegans
Species Accounts
1. Cyrtostachys bakeri Heatubun sp. nov. palma montana elata caespitosa 3 truncis, corona 8 foliis porrectis,
petiolo breve, 4 – 10 cm longo, foliolis pendulis, crassis
coriaceis, in sicco supra griseo-brunneis, infra palidioribus, inflorescentia in 3 ordines ramificanti, staminibus
12, pistillodio 3 lobis teretis membranaceis, distincta.
Typus: Papua New Guinea, Western Province, North Fly
Distr., Junction of Harvey Creek and Ok Mani R., 10 km
WNW of Tabubil, Dec. 2000, Baker et al.1138 (holotypus
K!; isotypi AAU, LAE, NY).
Robust, clustering tree palm 15 – 25 m, with up to 3 – 7
adult stems. Stem 15 – 25 cm in diam.; internodes 10 –
20 cm long; crown appearing shuttle-cock shaped.
Leaves 8 in crown, almost erect, with pendulous leaflets,
400 – 450 cm long (including petiole); sheath tubular,
110 – 130 × 40 cm; crownshaft c. 180 cm long, 15 –
25 cm in diam., whitish to mid-green with numerous
fine purple scales; petiole 4 – 10 cm long, c. 4 cm wide
and 2 cm thick at the base, channelled adaxially,
rounded abaxially; leaflets regularly arranged, leathery,
78 – 90 leaflets on each side, middle leaflets c. 92 ×
5 cm, apical leaflets c. 30 × 1.5 cm, briefly pointed and
sometimes notched at apices, green, discolorous when
dried, brown adaxially, paler abaxially, with fine sparse
ramenta on mid-vein on abaxial surface. Inflorescence
infrafoliar, strongly divaricate, 90 – 120 cm long,
branched to 3 orders, c. 15 primary branches, proximal
pair strongly recurving; prophyll and bract caducous,
not seen; peduncle c. 16 cm, caducous, not seen;
rachillae c. 55 – 57 cm long and c. 9 mm diam., rachilla
bract conspicuous, low, calyx persistent on rachillae
after fruits fallen; 20 – 22 pits in 1 cm of rachilla length
(at fruiting stage), pits 2 mm in diam. Staminate flowers c.
2.3 × 2.3 mm, asymmetrical; sepals c. 2 × 2 mm,
imbricate, rounded, strongly keeled; petals c. 2 ×
1.75 mm, triangular, brown in apical and basal parts;
stamens 12; filaments c. 1 × 0.1 mm; anthers 1 – 1.1 ×
0.5 – 0.7 mm; pollen size, long axis 34 – 45 µm, short
axis 35 – 40 µm, proximal wall thickness 2 – 3 µm,
distal wall thickness 2 – 3 µm, tectum surface verrucate,
trichotomosulcate grains present; pistillode 0.8 ×
0.3 mm, trifid, separated almost to the base, membranous. Pistillate flowers c. 3 × 2.8 mm; sepals 2 – 3 × 2 –
2.5 mm, imbricate, strongly keeled and dark brown to
black; petals 2 – 2.3 × 1.5 – 2 mm; gynoecium 1.8 ×
0.8 mm (including stigma 0.5 mm); staminodes circular, membranous. Fruits 9 – 25 × 3 – 5 mm, elongated,
pointed tips, hard, black when ripe. Seeds 7 – 8 × 4 –
4.5 mm, ovoid, rounded apically and flattened basally,
hard; endosperm homogenous. (Figs. 2A – C, 5).
DISTRIBUTION. Known only from the type collection
from Tabubil area in North Fly Distr., Western
Province in Papua New Guinea and one collection in
cultivation in Lae Botanic Garden.
SPECIMENS EXAMINED. PAPUA NEW GUINEA. Western
Province, North Fly Distr., Junction of Harvey Creek
and Ok Mani R., 10 km WNW of Tabubil, c. 750 m a.s.l.,
Dec. 2000, Baker et al. 1138 (holotype K!; isotypes AAU,
LAE, NY). CULTIVATED. Lae Botanic Garden, main palm
collection, Nov. 2006, Baker & Fazang 1308 (K, LAE,
MAN).
HABITAT. Growing in the forest on valley side near the
river margin at an altitude of c. 750 m above sea level.
VERNACULAR NAME. Not known.
USES. Not known.
CONSERVATION STATUS. Data Deficient (DD). The
conservation status of this palm is not known. The
information about this species is based on just one
collection from the vicinity of Tabubil area, Papua
New Guinea. More population and distribution data
are required.
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80
KEW BULLETIN VOL. 64(1)
Fig. 5. Cyrtostachys bakeri. A habit; B apical portion of leaf; C middle portion of leaf; D leaf sheath, upper part; E portion of
inflorescence; F detail of rachilla; G portion of rachilla with perianth of pistillate flowers persisting; H, J staminate flower whole and
in section; K pistillate flower. Scale bar: A = 5 m; B, C = 8 cm; D = 20 cm; E = 6 cm; F, G = 1 cm; H, J = 1.6 mm; K = 3 mm. All
from Baker 1138. DRAWN BY LUCY T. SMITH.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
NOTES. Cyrtostachys bakeri differs from all other species
in its leathery leaflets that are relatively widely spaced,
in its pits being the most congested in the genus with
20 – 22 pits in a 1 cm length of rachilla, and in its
membranous pistillode with lobes separate to the base.
It is also the only species that possesses both a shuttlecock shaped crown and pendulous leaflets. It is most
similar to C. elegans in the robust, clustering habit and
pendulous leaflets, but differs in the shuttle-cockshaped crown, the inflorescence branched to 3 orders,
the 12 stamens and the habitat at relatively high
elevation (c. 750 m a.s.l.).
Cyrtostachys bakeri is named for the collector of the
specimen, Dr William J. Baker, palm botanist and head
of the palm section of the Royal Botanic Gardens,
Kew, UK, in recognition of his contributions to palm
botany as a whole and in New Guinea, in particular.
2. Cyrtostachys barbata Heatubun sp. nov. palma
solitaria montana, habitu C. loriae Becc. similis, sed
petiolo 16 – 20 cm longo, petiolo rachideque indumento pallido vel brunneo dense tecto, foliolis confertis, crassis, in sicco supra glaucis vel brunneis, infra
griseis, infra secus costam ramentis brunneis dense
tectis, inflorescentia parva rachillis gracilibus foveis
vadosis ferentibus, staminibus 8 – 10, differt. Typus:
Indonesia, Papua Province, Sarmi distr., Mamberamo,
Idenburg R., 4 km of SW Bernard Camp, 900 m a.s.l.,
March 1939, Brass 13707 (holotypus L!; isotypi A!, BO!)
Large, solitary tree palm to 25 m. Stem 13 – 20 cm in
diam., glossy green apically and brown basally, nodal
scars very conspicuous. Leaves c. 348 cm long (including
petiole); sheath tubular, c. 143 cm long, c. 38 cm diam.,
fibrous at margin near petiole, forming distinct crownshaft, green; petiole 16 – 20 cm long, 4 – 5 cm wide and
1.5 – 2 cm thick at the base, covered in thick white
indumentum, and scattered filamentous, brown scales;
leaflets regularly arranged, pectinate, leathery, number
of leaflets on each side not known, middle leaflets c.
105.5 × 6.5 cm, apical leaflets c. 23 × 2.5 cm, briefly
pointed with long tip and sometimes rounded and
notched at apices, green, discolourous when dried, light
brown to glaucous adaxially, paler brown to whitish
abaxially, with thick, membranous brown ramenta,
inflexed, beard-like and continuous along mid-vein
abaxially. Inflorescence slender, branched to 3 orders, light
brown to whitish brown when dried; rachilla 43 – 49 cm
long; c. 4 mm diam.; pits superficial to shallow, 13 pits in
1 cm length rachilla (at fruiting stage), pit 2 – 3 mm
diam. Staminate flowers c. 3.5 × 2.6 mm; sepals 1.5 – 2 ×
2 mm; petals c. 2.5 × 1.8 mm; stamens 8 – 10; filaments
2.5 × 0.2 – 0.5 mm; anthers 1 – 1.1 × 0.5 – 0.8 mm;
pollen size long axis 38 – 53 µm, short axis 24 – 35 µm,
proximal wall thickness 3 – 4 µm, distal wall thickness 1 –
3 µm, tectum surface gemmate, trichotomosulcate grains
81
present; pistillode 1.5 × 0.5 – 0.8 mm. Pistillate flowers c.
3 × 3.3 mm; sepals 2.5 – 2.7 × 2.5 – 3 mm; petals 3.5 –
4 × 3 – 4 mm; gynoecium c. 2.8 × 1.2 mm (including
stigma). Fruits c. 12 × 5 mm, ellipsoid to sickle-shaped;
beak 2 mm long. Seeds c. 5 × 4 × 4 mm, ovoid, rounded
apical and flatted basally. (Figs. 2D – E, 6).
DISTRIBUTION. This palm is known only from its type
locality in the Idenburg R. area, Mamberamo basin,
Indonesian Province of Papua.
SPECIMEN EXAMINED. INDONESIA. Papua Province.
Sarmi distr.; Mamberamo, Idenburg R., 4 km of SW
Bernard Camp, 900 m a.s.l., March 1939, Brass 13707
(holotype L!; isotypes A!, BO!)
HABITAT. Growing in primary forest on slopes in
Agathis forest at an altitude of 900 m above sea level.
VERNACULAR NAME. Nibung (Indonesian dialect in
Papua, also used for other tree palms).
USES. Not known.
CONSERVATION STATUS. Data Deficient (DD). Cyrtostachys barbata is only known from one collection made
by L. J. Brass in Mamberamo basin, Indonesian
Province of Papua, although in Brass’s field notes he
noted that the palm was common in Agathis forest and
rain forest on slopes. Agathis is an important timber
tree in New Guinea and provides a luxury wood for
the timber trade, so the extensive logging activities in
that area will very likely affect the entire population of
this palm. More populations and distribution data are
required to assess the conservation status of this species.
NOTES. Cyrtostachys barbata is similar to C. loriae in habit,
but differs from the latter and other species in the
presence of continuous beard-like brown ramenta
along the mid-veins on the abaxial surface of leaflets,
the very short space between leaflets giving the leaf a
pectinate appearance, rachis and petiole covered by
thick white indumentum and scattered filamentous
brown scales; the slender rachillae with congested
superficial pits and the habitat in Agathis forest at high
altitude (900 m a.s.l.).
Cyrtostachys barbata also differs from C. elegans in its
solitary habit, the elongate petiole and inflorescences
branched to three orders, rather than clustering habit,
short petiole and inflorescence branched to four
orders. It also differs from C. glauca in its solitary
habit and hemispherical crown outline, rather than
clustering habit and spherical crown.
Unlike other species in the genus, Cyrtostachys barbata
and C. glauca have slender rachillae with superficial to
shallow pits, but C. barbata has brown rachillae when
dried and congested pits c. 13 per 1 cm rachilla length,
while C. glauca has white rachillae when dried and
sparse pits 3 – 5 per 1 cm rachilla length.
The specific epithet came from Latin word barbatus
meaning bearded, in reference to the dense ramenta
along the mid-vein that contrast with the white colour
of the abaxial leaflet surface. This specimen was seen
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
82
KEW BULLETIN VOL. 64(1)
Fig. 6. Cyrtostachys barbata. A apical portion of leaf; B middle portion of leaf; C detail of ramenta; D portion of inflorescence; E
portion of rachilla in fruit; F, G staminate flower whole and in section; H, J pistillate flower whole and in section. Scale bar: A, B =
8 cm; C = 5 mm, D = 6 cm; E = 1 cm; F – J = 2.5 mm. All from Brass 13707. DRAWN BY LUCY T. SMITH.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
by H. E. Moore in 1956 and identified as Cyrtostachys
ledermanniana Becc. with a question mark behind that
name. A few characters of C. barbata, especially regarding leaf and rachilla, fit with C. ledermanniana based
on Beccari’s description in the protologue (Beccari
1923). However, C. ledermanniana was described from
very inadequate material; Beccari made his description based on two different Ledermann collections
(Ledermann 9063 and 7175). These specimens came
from two different areas in Papua New Guinea.
Ledermann 9063 was collected from Ettapenberg at
850 m a.s.l., and from this specimen, Beccari made a
brief description of habit, leaves, inflorescence,
flowers, fruits and seeds. He wrote his description of
the staminate flowers from Ledermann 7175 from the
Sepik area in the swamp forest at an altitude 20 –
40 m a.s.l. The type of C. ledermanniana, and almost
all the Ledermann’s collections were destroyed
during the Second World War in Berlin, and we have
been unable to locate another duplicate. Thus, C.
ledermanniana is treated as an imperfectly known
taxon in this monograph.
3. Cyrtostachys elegans Burret (1937: 472). Type:
Cultivated in Bogor Botanic Garden, II.F. 17. ex New
Guinea, April – May 1936, Furtado SFN 3/1/28
(holotype B†; isotype K!, L!, SING).
Robust, clustering tree palm to 15 (– 20) m, with up to c.
3 adult stems and 4 – 6 or more suckers at the base,
crown hemispherical in outline. Stem 15 – 30 cm diam.,
greyish green apically, greyish brown to black near base;
internodes c. 25 cm long. Leaves 9 in crown, curved, 300
– 350 cm long (including petiole); sheath tubular, c.
33 cm wide, forming a distinct crownshaft, c. 250 cm
long, light green to pale yellow; petiole short to 10 cm
long, 3.5 – 4 cm wide and 1.5 – 2.5 cm thick at the base,
channelled adaxially, rounded abaxially; leaflets pendulous, regularly arranged, leathery, 100 – 102 leaflets on
each side, middle leaflets 100 – 126.5 × 3.5 – 4.5 cm,
apical leaflets c. 33.5 × 1 cm, briefly pointed and
sometimes notched at apices, green, discolorous when
dried, light brown adaxially, pale brown to whitish
abaxially, fine brown ramenta discontinuous along midvein on abaxial surface. Inflorescence 75 – 100 cm long, c.
160 cm wide, branched to 4 orders, creamy to yellowish
green, brown when dried; prophyll c. 65 × 24 cm,
leathery, peduncular bract similar to prophyll; peduncle very short to 7.5 cm; rachillae 34 – 61.5 cm long;
17 – 19 pits per 1 cm rachilla length (at fruiting stage),
pits 2 – 4 mm diam. Staminate flowers 2 – 2.5 × 2 –
2.7 mm, asymmetrical; sepals 1.8 – 2.3 × 1.5 – 2 mm;
petals 1.4 – 2 × 1.3 – 1.5 mm; stamens 9; filaments 0.5 –
1.5 × 0.1 – 0.3 mm; anthers 0.5 – 1 × 0.3 – 0.7 mm;
pollen size long axis 31 – 42 µm, short axis 24 – 42 µm,
proximal wall thickness 1 – 3 µm, distal wall thickness
83
1 – 2 µm, tectum surface verrucate, less frequently
gemmate, trichotomosulcate grains present; pistillode
0.8 – 1.3 × 0.3 – 0.5 mm, trifid. Pistillate flowers 2 – 5.2 ×
1.3 – 4.5 mm; sepals 1.6 – 4.8 × 1 – 3.5 mm; petals 1.2 –
3.8 × 0.5 – 3.9 mm; gynoecium 0.9 – 4.5 × 0.4 – 2.5 mm
(including stigma); staminodes 4 – 5, membranous.
Fruits 12 – 17 × 5 – 6 mm, ellipsoid to sickle-shaped,
green to black (when mature); beak 1 – 2 mm long,
perianth persistent, forming a narrow cylinder at the
base or leaving a different coloured scar, ⅓ – ½ length
of fruit. Seeds 6 – 7 × 4 – 5 × 4 mm, ellipsoid, rounded
apical and flatted basally. (Fig. 2F – H).
DISTRIBUTION. Cyrtostachys elegans is a Central-West New
Guinean species and is known only from lowlands in
Nabire and Timika in the Indonesian Province of
Papua.
SPECIMENS EXAMINED. INDONESIA. Papua Province,
Nabire distr.; Wanggar, Bumi R., Satuan Pemukiman
Transmigrasi C (SP-C), c. 10 m a.s.l., Feb. 2001,
Heatubun et al. 341 (K!, AAU!, MAN!); Bivak Kanehira,
10 km inward from mouth of Bumi R., c. 100 m a.s.l.,
March 1940, Kanehira & Hatusima 12851 (A!); Bumi
R., 40 km inward of Nabire, c. 300 m a.s.l., March
1940, Kanehira & Hatusima 12747 (A!). Mimika distr.,
Timika area, PT. Freeport Indonesia Concession area,
East levee by drowned forest, 5 km S of Kampung Kali
Kopi, alt. 20 m a.s.l., Feb. 1998, Heatubun et al. 194
(AAU!, BH!, BO!, K!, L!, MAN!). CULTIVATED. Bogor
Botanic Garden origin from New Guinea, II.F.17, May
1936, Furtado SFN 3/1/28 (K!, L!, SING isotypes); XII.
E.40, April – May 1936, Furtado SFN 3/1/06 (K!, L!,
SING).
HABITAT. This palm grows in swampy areas in lowland
rain forest at an altitude 10 – 300 m above sea level.
VERNACULAR NAME. Nibung (Indonesian dialect in
Papua, also used for other tree palms).
USES. Not known.
CONSERVATION STATUS. Near Threatened (NT). The
resettlement and relocation of Nabire town after being
hit by a large earthquake recently, and the development in the Timika area to support PT. Freeport
Indonesia (the world biggest copper and gold mining
company) mining activities will affect the population
of this palm. Large areas of forest in the lowlands will
disappear.
NOTES. Cyrtostachys elegans is similar to C. loriae in being
a robust tree palm with pendulous leaflets and short
petiole, but differs from the latter in its clustering
habit, the hemispherical crown and elongate inflorescence branched to 4 orders, rather than solitary habit,
spherical crown and robust inflorescence branched to
3 orders. This species also differs from the other New
Guinean robust tree palm C. bakeri in the presence of
curved leaves and hemispherical crown, rather than
erect leaves and shuttle-cock crown; however, both
species have pendulous leaflets.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
84
This species was described by Burret in 1937 based
on a herbarium specimen made by C. X. Furtado from
a plant cultivated in the Bogor Botanical Garden.
4. Cyrtostachys excelsa Heatubun sp. nov. palma
solitaria calciphila, corona 7 foliis erectis rigidisque,
petiolo breve, 11 – 15 cm longo, foliolis irregulariter
binatim dispositis, crassis coriaceis, in sicco supra atrobrunneis, infra palidioribus, inflorescentia in 4 ordines
ramificanti, rosea vel purpurea, staminibus 12 differt.
Typus: Indonesia, Papua Province, Nabire distr., Kwatisore, Pintu Angin, km-14 road of PT. Kaltim Hutama,
Feb. 2001, Heatubun et al. 330 (holotypus K!; isotypi
AAU!, MAN!).
Robust, solitary tree palm to 25 m. Stem c. 15 cm diam.,
greyish to white in colour, nodal scars conspicuous;
internodes c. 30 cm long, crown appearing shuttle-cockshaped in outline. Leaves c. 7 in crown, stiff, slightly
recurved, 250 – 300 cm long (including petiole); sheath
tubular, c. 30 cm wide, forming a distinct crownshaft;
petiole 11 – 15 cm long, c. 4 cm wide and 2 cm thick at
the base, channelled adaxially, rounded abaxially, with
thin, caducous, dark brown indumentum; leaflets
irregularly arranged in 1s or 2s, leathery, c. 80 leaflets
on each side, middle leaflets c. 112 × 4 cm, apical
leaflets c. 28.5 × 1.7 cm, briefly pointed and sometimes
notched at apices, green, concolorous when dried, light
to chocolate-brown on both surfaces; rachis with thin,
caducous, fine lepidote indumentum, interspersed with
white scales. Inflorescence infrafoliar, strongly divaricate,
c. 100 cm long, branched to 4 orders, pink to reddishpurple when young to greenish-grey when mature and
dark brown when dried; peduncle very short; rachilla
50 – 75 cm long; c. 15 pits per 1 cm rachilla length
(before anthesis). Staminate flowers 2.5 × 2.5 mm,
asymmetrical; sepals c. 2.1 × 2 mm, imbricate, rounded,
strongly keeled; petals c. 1.9 × 1.8 mm, striate,
triangular; stamens 12; filaments 1 × 0.1 mm; anthers
1 × 0.5 – 0.6 mm; pollen size: long axis 35 – 42 µm,
short axis 33 – 42 µm, proximal and distal wall
thickness 2 – 3 µm, tectum surface verrucate, less
frequently gemmate, trichotomosulcate grains present;
pistillode 1 × 0.3 mm, trifid. Pistillate flowers c. 3.1 ×
2.5 mm; sepals c. 2.2 × 2 mm, imbricate, strongly
keeled, dark brown to black; petals c. 2.5 × 1.5 mm,
triangular; gynoecium 1.8 × 0.5 mm (including stigma
0.7 mm); staminodes 3, triangular, membranous. Fruits
not seen. Seeds not seen. (Fig. 2J – L).
DISTRIBUTION. Only known from one locality in forest
on slopes at Pintu Angin (14 km on the road of the
logging company PT. Kaltim Hutama) near Kwatisore
village, Nabire distr., Papua.
SPECIMEN EXAMINED. INDONESIA. Papua Province.
Nabire distr.; Kwatisore, Pintu Angin, km-14 road of
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
KEW BULLETIN VOL. 64(1)
PT. Kaltim Hutama, c. 500 m a.s.l., Feb. 2001,
Heatubun et al. 330 (holotype K!; isotypes AAU!,
MAN!).
HABITAT. This palm grows in limestone hill forest at an
altitude c. 500 m above sea level.
VERNACULAR NAME. Warita (Jamur dialect); Mojigre
(Kwatisore dialect); Nibung (Indonesian dialect in
Papua, also used for other tree palms).
USES. Stems and leaves are used as materials to
construct traditional houses (flooring and thatch).
CONSERVATION STATUS. Endangered (EN B2a – c).
Cyrtostachys excelsa meets criteria B2a – c for threat
category “Endangered” (IUCN 2001) because its area
of occupancy is less than 500 km2 occurring only on a
limestone hill in Kwatisore, Nabire, Indonesian
Province of Papua. Furthermore, forest in that area
was heavily disturbed by logging activities.
NOTES. Cyrtostachys excelsa is the only species in the
genus with grouped leaflets (irregularly arranged
leaflets), concolorous when dried. This species, together with C. elegans, has inflorescences branched to
4 orders, the highest number of branching orders in
the genus — the other species being branched to 2
and 3 orders. Also C. excelsa is the only species from
the Papuasian region that has pink to reddish purple
inflorescences, like C. renda from west Malesia.
The crown of Cyrtostachys excelsa is quite distinct,
being composed of a small number of leaves (c. 7),
forming a shuttle-cock shape and porrect leaflets, with
flattened lowest leaf and other leaves slightly recurved
near the tips.
The specific epithet for this species came from the
Latin word excelsus which means lofty or high, reflecting the impression of the palm when first seen by the
author.
5. Cyrtostachys glauca H. E. Moore (1966: 86). Type:
Papua New Guinea, Morobe Province, Morobe district, forested slopes along logging road just beyond
bridge over Markham R. on the road to Bulolo, March
1964, Moore 9272 (holotype BH!; isotype LAE).
Slender to moderate, clustering tree palm to 5.5 – 15
(– 21.6) m, with up to c. 3 adult stems and up to 4 or more
suckers at base, crown spherical in outline. Stem 5 – 15 cm
diam., bright green and becoming grey in age, nodal
scars conspicuous; internodes 3.5 – 7.5 cm long. Leaves
6 – 8 in crown, leaves spreading, 172 – 410 cm long
(including petiole); sheath tubular, 66 – 130 cm long,
8 – 22 cm wide, forming a distinct crownshaft, 70 –
240 cm long, bright glaucous, sparsely and minutely
lepidote with membranous brown scales; petiole green,
elongate, 25 – 88 (– 100) cm long, 2 – 3 cm wide and
1 – 1.6 cm thick at the base, rounded and rather densely
and minutely brown lepidote below, concave and
similarly lepidote or puncticulate above; leaflets regularly
85
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
arranged, papery, 45 – 67 leaflets on each side, middle
leaflets 58 – 99 × 3 – 5 cm, apical leaflets 12. 5 – 24 ×
0.5 – 2.5 cm, briefly pointed and sometimes notched
at apices, green, discolorous when dried, glaucous
adaxially, glaucous to whitish abaxially, with thick, brown
membranous ramenta, discontinuous along mid-vein on
abaxial surface; rachis with similar lepidote indumentum
as petiole. Inflorescence infrafoliar, strongly divaricate, 50 –
130 cm long, 100 – 200 cm wide, branched 3 – 4 (mostly
3) orders, green to pale yellow in colour and pale brown
to whitish-green when dried; peduncle very short to
5 cm; rachillae 25 – 50 cm long; 2 – 4 mm diam., white,
calyx persistent when fruits fallen; pits superficial to
shallow, 3 – 5 pits per 1 cm rachilla length, pit 2 – 3 mm
diam. Staminate flowers 2.5 – 3.2 × 2.5 – 2.8 mm; sepals
2 – 2.5 × 2 – 2.3 mm; petals 2.2 – 2.9 × 1.8 – 2.1 mm;
stamens 9 – 10; filaments 0.9 – 2 × 0.1 – 0.3 mm; anthers
1.3 – 1.5 × 0.5 – 0.8 mm; pollen size, long axis 35 –
45 µm, short axis 23 – 41 µm, proximal wall thickness
3 µm, distal wall thickness 3 – 4 µm, tectum surface
verrucate, trichotomosulcate grains absent; pistillode
1.1 – 1.8 × 0.4 – 0.5 (1 – 1.5 at the base) mm. Pistillate
flowers 3 – 3.5 × 2.5 – 3 mm; sepals 2.8 – 3 × 2.8 mm;
petals 2.7 – 3.1 × 2.5 – 3 mm; gynoecium 1.5 – 2.5 ×
0.5 – 1.5 mm (including 3 recurved stigmas); staminodes
3, membranous. Fruits 9 – 12 × 4 – 6 mm, ellipsoid to
sickle-shaped, green to black; beak 1 – 4 mm long. Seeds
5 – 7 × 3 – 4 × 3 – 4 mm, ellipsoid, rounded apical and
flattened basally. (Figs. 3A – C, 7).
DISTRIBUTION. Cyrtostachys glauca is a Papua New
Guinean species. This palm is distributed in Morobe,
Milne Bay and Central Provinces.
SPECIMENS EXAMINED. PAPUA NEW GUINEA. Central
Province; Yamu road, c. 15 km NE of Cape Rodney,
30 m a.s.l., Sept. 1969, Pullen 8212 (L!, LAE); Pullen
8231 (BH!, CANB, L!, LAE); Yamu village near Mori
R., Sept. 1969, Pullen 8198 (BH, CANB, L!, LAE,).
Milne Bay Province; along Kabawawa highway, 0 m
a.s.l., March 2000, Barfod et al. 454 (AAU!, BRI,
CANB, K!, LAE); Esa’ala, Ferguson Island, E of Lake
Lavu, 100 m a.s.l., Nov. 1976, Croft et al. LAE 68805
(BRI, L!, LAE); Maiyu R., c. 16 km WNW of Biniguni
airstrip, c. 280 m a.s.l., June 1972, Pullen 8409 (A,
CANB, L!, LAE,). Morobe Province; forested slopes
along logging road just beyond bridge over Markham
R. on the road from Lae to Bulolo, March 1964, Moore
9272 (holotype BH!); Lae, Garagos logging road,
Katik LAE 62219 (BRI, L!, LAE); Bulili Ridge, near
Lababia guest house, 50 – 400 m a.s.l., Kjaer & Magun
512 (AAU!, BRI, CANB, K!); just before Gwabadik on
the 4WD track, 275 m a.s.l., Feb. 1993, Takeuchi 8770
(A!, L!, LAE); NW of Waria R., Buttress Ridge above
Wara Pao, near Yai Village, c. 200 m a.s.l., July 1999,
Takeuchi et al. 13217 (A!); Lae, 60 m a.s.l., Sept.1971,
Essig & Katik LAE 55009 (CANB, L!, LAE,); LAE
62219 (L!, LAE).
CULTIVATED. Lae Botanic Garden, main palm collec-
tion, Nov. 2006, Baker & Fazang 1309 (K, LAE).
HABITAT. Growing in primary forest or secondary forest
with scattered subsistence gardens in lowlands to
sloping terrain or hill forest at an altitude 30 –
400 m above sea level.
VERNACULAR NAME . Vekintambu (Lababia dialect;
Morobe), Hek (Madang dialect).
USES. Stem for building purposes.
CONSERVATION STATUS. Data Deficient (DD). Although
this palm has a relatively wide distribution in SE Papua
New Guinea, suggesting the category “Least Concern”,
more accurate data related to distribution of, and the
threats to, this species are still needed to assess its
conservation status.
NOTES. Cyrtostachys glauca is easily distinguished from
all other species by its spherical crown, glaucous leaf
sheath and crownshaft, slender and elongate petioles
of more than 50 cm long (– 100 cm), papery leaflets,
slender inflorescence and rachilla with sparse superficial
pits, and by the unique presence of trichotomosulcate
pollen grains.
This is a very peculiar species within Cyrtostachys
because of the nature of the pits along the rachilla.
The pits are very shallow or even lacking in a few
specimens (Kjaer & Magun 512 and Barfod 454) and
together with the tiny flowers (in bud) the species may be
superficially confused with Heterospathe or Rhopaloblaste.
However, the generic characters such as presence of a
crownshaft, connate petals and stamens in staminate
flowers, and fruit and seed morphology still clearly
indicate it as a species of Cyrtostachys.
6. Cyrtostachys loriae Becc. (1905: 303). Type: Papua
New Guinea, Owen Stanley Range Province, 1887,
Hartmann s.n. (holotype FI!).
Cyrtostachys kisu Becc. (1914a: 289); synon. nov. Type:
The Solomon Islands, Faro island, Guppy 235
(holotype K!).
Cyrtostachys peekeliana Becc. (1914b: 28); synon. nov.
Type: Papua New Guinea, New Ireland, Peekel 106
(holotype B†; isotype FI!).
Cyrtostachys brassii Burret (1935: 328); synon. nov.
Type: Papua New Guinea, Central Province, Kubuna,
Nov. 1933, Brass 5600 (holotype B†; isotype A!).
Cyrtostachys phanerolepis Burret (1936: 324); synon. nov.
Type: Papua New Guinea, Morobe, Sattleberg, July
1935, Clemens 1353 (holotype B†; isotype L!, K,
photo!).
Cyrtostachys microcarpa Burret (1939: 203); synon. nov.
Type: Papua New Guinea, Western Province, Fly R.,
Lake Daviumbu, Sept. 1936, Brass 7757 (holotype A;
isotype L!).
Robust, solitary tree palm to 10 – 30 m. Stem 11.5 –
30 cm diam., brownish-grey to whitish below and
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
86
KEW BULLETIN VOL. 64(1)
Fig. 7. Cyrtostachys glauca. A apical portion of leaf; B middle portion of leaf; C portion of inflorescence; D detail of rachilla with
flowers in bud; E portion of rachilla in fruit; F, G staminate flower whole and in section; H, J pistillate flower whole and in section; K
fruit in section. Scale bar: A, B = 8 cm; C = 6 cm; D, K = 5 mm; E = 1 cm; F, G = 2 mm; H, J = 2.5 mm. A – B, H – J from Kjaer 512, C –
D, F – G, K from Moore 9272. DRAWN BY LUCY T. SMITH.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
green to olive-green above, nodal scars conspicuous,
internodes 3 – 30 cm long, crown hemispherical in
outline. Leaves 8 – 14 in crown, leaves spreading, 250 –
480 cm long (including petiole); sheath tubular,
112.5 – 180 cm long, 25 – 68 cm wide, forming
distinct crownshaft, 125 – 200 cm long, pale yellow to
light green; petiole almost missing to short (1 – 10 cm
long), 3 – 5.5 cm wide and 1 – 2.8 cm thick at the
base, with thin or thick brown to whitish-purple
lepidote indumentum; rachis with similar indumentum
as the petiole; leaflets regularly arranged, leathery, 76 –
189 leaflets on each side, middle leaflets 80 – 152 × 3.4 –
6.8 cm, apical leaflets 14 – 50 × 0.8 – 2 cm, briefly
pointed and sometimes notched at apices, green,
discolorous when dried, glaucous adaxially, glaucous to
whitish abaxially, fine brown ramenta discontinuous
along mid-vein on abaxial surface. Inflorescence infrafoliar, strongly divaricate, 43 – 150 cm long, up to
250 cm wide, branched to 3 orders, green to pale
yellow, light brown to black when dried; peduncle very
short to 10 cm; rachillae 25 – 88.5 cm long; 6 – 9 mm
diam., brown to rusty brown, calyx persistent after fruits
fallen; 8 – 16 pits per 1 cm rachilla length (in various
stages), pits 2 – 6 mm in diam., deep. Staminate flowers
2.5 – 4 × 2 – 3 mm; sepals 1.2 – 2.2 × 1.5 – 2.9 mm;
petals 2 – 3 × 1.2 – 2.5 mm; stamens 9 – 13; filaments
0.7 – 3.5 × 0.1 – 0.2 mm; anthers 1 – 1.5 × 0.5 –
0.8 mm; pollen size, long axis 27 – 56 µm, short axis
25 – 48 µm, proximal wall thickness 1.5 – 5 µm, distal
wall thickness 1 – 5 µm, tectum surface sparsely
verrucate and gemmate, trichotomosulcate grains
present; pistillode 0.7 – 1.7 × 0.2 – 1 (1 – 1.5 at the
base) mm. Pistillate flowers 2 – 6 × 2.2 – 5.5 mm; sepals
2.5 – 5.2 × 2.1 – 6.2 mm; petals 1.5 – 5.2 × 1 – 5 mm;
gynoecium 1 – 5 × 0.5 – 3.5 mm (including 3 recurved
stigmas); staminodes triangular (3 – 4) to circular,
membranous. Fruits 8 – 16 × 4 – 5 mm, ellipsoid to
sickle-shaped, green to black; beak 0 – 2 mm long. Seeds
5 – 8 × 2 – 5 × 2 – 5 mm, ellipsoid to ovoid, rounded
apically and flattened basally. (Figs. 3D – H, 8).
DISTRIBUTION. Cyrtostachys loriae is a widespread species
in the Papuasian region, distributed from Kepala
Burung (Bird Head’s Peninsula) in the west to
Solomon Islands in the east.
SPECIMENS EXAMINED. INDONESIA. West Papua Province:
Manokwari distr.; Ransiki subdistr., Ransiki, Kostermans
s.n. (BO!). Papua Province: Yapen distr.; Yapen Island,
Yapen Tengah Mts. Natural Reserves, near base camp
at Mananayang R., c. 500 m a.s.l., Feb. 2005, Heatubun
et al. 546 (BO!, K!, MAN!); Trans Yapen road, first
slope from Mananayang R. to Serui, c. 550 m a.s.l.,
Feb. 2005, Heatubun et al. 547 (BO!, K!, MAN!).
Mimika distr., Timika, mile 50 on road to Tembagapura, forest to W of container depot, 545 m a.s.l., Feb.
1998, Heatubun et al. 208 (BH!, BO!, K!, L!, MAN!).
Sarmi distr., Mamberamo R., Idenburgh R., Bernard
87
Camp, 55 m a.s.l., April 1939, Brass 13807 (A!, L!);
Bernard Bivak (Camp), 50 m a.s.l., Meijer Dress 501
(BO!, L!). Jayapura distr., Cyclops Mts, N Cyclops Mts.
Natural Reserves, 50 – 700 m a.s.l., Jan. 2001, Desianto
01 (AAU, K!, MAN!); foot path to Raveni summit,
c. 790 m a.s.l., Aug. 1998, Heatubun et al. 279 (BO!,
FTG!, K!, L!, MAN!). Keerom distr., Arso, Tami R.,
Treferer, 100 – 150 m a.s.l., March 2002, Gusbager et al.
23 (K!, LAE!, MAN!); Workwana village, Yumseyus,
50 – 100 m a.s.l., Feb. 2005, Heatubun et al. 527 (BO!,
K!, MAN!); Heatubun et al. 532 (BO!, K!, MAN!);
Heatubun et al. 533 (BO!, K!, MAN!). Merauke distr.,
Kwell village, 60 m a.s.l., Sept. 2000, Maturbongs 654
(BO!, AAU, K!, MAN!); Bot R., about half-way between
Bupul and Lake Wam, 60 m a.s.l., Aug. 1954, van Royen
4734 (A!, BO!, L!). PAPUA NEW GUINEA. Sandaun
Province: Miwaute, Nov. 1996, Barfod et al. 395
(AAU!). Western Province: Palmer R., 2 miles below
junction Black R., July 1936, Brass 7162 (A!, L!);
Middle Fly R., Lake Daviumbu, Sept. 1936, Brass
7757 (A, L!); c. 2 miles N of Kiunga, Upper Fly R.,
c. 300 m a.s.l., Sept. 1967, Pullen 7305 (BH, CANB, L!,
LAE,). East Sepik Province: Sepik distr.; along the
Sepik R. between Ambunti and Malu, 50 m a.s.l., May
1966, Hoogland & Craven 10114 (A!, BH, CANB, K!, L!,
LAE); Anggoram subdistr.; Wewak – Anggoram area,
4 miles N of Timbunke mission, 30 m a.s.l., Sept. 1959,
Pullen 1692 (CANB, K!, L!, LAE). West Sepik Province:
Sepik distr., Aitape subdistr.; along Pieni R. near
Walwali Village, 30 m a.s.l., June 1961, Derbyshire &
Hoogland 8020 (A, K!, L!). Telefomin distr.; Carpentaria Exploration Co. Frieda R. Camp, 60 m a.s.l., April
1978, Essig & Young LAE 74052 (CANB, L!, LAE); Hak
Valley, c. 900 m a.s.l., Morren & Frodin 3189 (K!).
Southern Highlands Province: Lake Kutubu, near Tage,
90 m a.s.l., Sept. 1961, Schodde 2248 (CANB, K!,). Gulf
Province: Kikori distr.; TFI logging concession, near
Morare village, 20 km NE of Kikori, 130 m a.s.l., Nov.
2000, Baker et al. 1110 (AAU, BRI, K!, L, LAE, NY);
logging camp on Vailala R., 0 m a.s.l., March 2000,
Barfod 478 (AAU!, BRI, K!, LAE); Barfod 482 (AAU!,
BRI, K!, LAE, CANB). Morobe Province: Sattleberg,
700 m a.s.l., July 1935, Clemens 1353 (K (photo)!, L!).
Milne Bay Province: Mullins bay road, 0 m a.s.l., March
2000, Barfod et al. 463 (AAU!, BRI, K!, LAE, CANB).
Central Province: Kuriva, 300 m a.s.l., March 2000,
Barfod 467 (AAU!, BRI, K!, LAE, CANB); Koitaki, 50 m
a.s.l., Carr 12253 (A, BM, L!); Kairuku subdistr.; c. 15
miles W of Maipa Village on Akaifu R., Sept. 1962,
Derbyshire 867 (CANB, L!, LAE). New Ireland Province:
New Ireland Island, Peekel 444 (FI!, K (photo)!). North
Solomon Province: Bougainville Island, Marmarromino,
50 m a.s.l., Oct. 1930, Kajewski 2220 (A!). THE SOLOMON
ISLANDS. Fauro island; Guppy 235 (holotype K!); Eastern
Peninsula; 200 m a.s.l., April 1964, Whitmore BSIP 3945
(BSIP, K!); New Georgia group; Baga Island, April 1964,
Whitmore BSIP 4210 (BSIP, K!).
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
88
KEW BULLETIN VOL. 64(1)
Fig. 8. Cyrtostachys loriae. A apical portion of leaf; B middle portion of leaf; C portion of inflorescence; D detail of rachilla; E
portion of rachilla in fruit; F, G staminate flower whole and in section. H, J pistillate flower whole and in section; K fruit in section.
A – B, from Baker 1110, C, D, H, J from Heatubun 208, E – G, K from Hoogland & Craven 10144. Scale bar: A, B, C = 8 cm; D, E =
1 cm; F, G = 2 mm; H, J = 3 mm; K = 7 mm. DRAWN BY LUCY T. SMITH.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
HABITAT. This palm grows in primary or secondary
forest in the lowlands or hill forest at an altitude of
30 – 400 m above sea level.
VERNACULAR NAMES. Papua Indonesian: tnang nyi
(Sentani, Jayapura); gap (Marap, Tami R.); terep/terrip
(Yei/Je, Merauke); Nibung (Indonesian dialect in
Papua, also used for other tree palms). Papua New
Guinea: yomberi (Timbunke, Sepik); yowoh (Waskuk,
Sepik); hek/he-ek (Amele, Madang); terep (Jal, Madang);
apaku (Mekeo, Maipa); flim (Mianmin); lobu (Wapi,
Marok); mun (Orme, Walwali); wai’eba (Kutubu);
toono-i (Bougainville Island); a ikul (New Ireland
Island). Solomon Islands: kwara’ae (Aatarae).
USES. Cyrtostachys loriae is one of the more useful species
in the genus from an ethnobotanical point of view. It has
been used traditionally by people native in both New
Guinea and the Solomon Islands. The stems and leaves
are used as building materials for traditional houses, e.g.
piles, flooring, water pipes, thatch and mattresses. The
palm heart or “cabbage” is also eaten fresh or cooked.
CONSERVATION STATUS. Least Concern (LC). Cyrtostachys
loriae is widespread in the Papuasian region, and as yet
the conservation status of this species seems not to be a
cause for concern. However, land conversion for oil
palm plantations or other purposes, including illegal
logging activities in West New Guinea (Indonesian
Provinces of Irian Jaya Barat and Papua) could have a
severe effect on the populations of the palm. Detailed
population studies are still needed to assess its conservation status more precisely.
NOTES. Cyrtostachys loriae was the first species of the
genus to be published from the Papuasian region
(Beccari 1905). This palm is easily distinguished by its
solitary and robust habit, spherical crown, pendulous
leaflets, very short (– 10 cm long) or missing petiole
and an inflorescence more robust than in other species,
branched to 3 orders, with robust rachillae bearing
large and deep pits.
Re-examination of the type specimens of Cyrtostachys
brassii, C. kisu, C. loriae, C. microcarpa, C. peekeliana and C.
phanerolepis revealed no significant differences among
them except those caused by differences in developmental stages, despite the inadequate nature of the
specimens. Morphological variation among them is continuous, especially after comparison with more adequate
specimens from recent collections. No disjunctions in
variation occur that would allow the consistent separation
of six species as recognised by previous authors. The
narrow species concept used in the past reflects limited
information obtained from single collections.
Moore (1966) pointed out his suspicions that the
five taxa above might not be distinct; they all have a
solitary habit and 12 stamens except for Cyrtostachys
phanerolepis — six stamens, pits in 9 series and larger
fruits (Burret 1936). Burret did not realise that the
specimen Clemens 1353 (the type of C. phanerolepis) had
been mixed with some male flowers of Licuala, and he
89
described C. phanerolepis with staminate flowers from
Licuala.
Cyrtostachys loriae is the most widespread species of
the genus in the Papuasian region, and also occupies a
wide range of ecological conditions from swampy
areas in the lowlands to heath forest in lower montane
vegetation, from evergreen rain forest to dry areas in
savannah lands and from the main island of New
Guinea to small off-shore islands and the Solomons.
The adaptation to various habitats is reflected in the
very variable appearance, a plasticity that occurs not
only in size and shape, but also the number of certain
organs, such as number of stamens. In some specimens, different numbers of stamens can be found
within one inflorescence or in different collections
from the same locality.
There are two collections from savannah areas in
Merauke, Indonesian Province of Papua, Maturbongs
654 and van Royen 4734, which look quite distinct in
the appearance of their leaves. The petiole and rachis
are covered by scaly indumentum, and the leaflets are
slender with discolorous surfaces (glaucous adaxially,
purplish-brown abaxially). Other characters such as
habit, stem, inflorescence and flowers, fruits and seeds
fit with Cyrtostachys loriae. The variation in leaf character may reflect different ecological conditions in
savannah areas.
7. Cyrtostachys renda Blume (1838: 66, 1843: 101).
Bentinckia renda (Blume) Mart. (1853: 316). Type:
Indonesia, Sumatra, East Sumatra, around Indrapura,
Korthals s.n. (lectotype L!, designated here; isolectotype K!).
Areca erythropoda Miq. (1861a: 6). Type: Indonesia,
Sumatra, Bangka Island, Djebus, Teisjmann s.n.
(holotype BO!).
Cyrtostachys lakka Becc. (1885: 141). Type: Malaysia,
Borneo, Sarawak, Kuching, Nov. 1866, Beccari PB
2674 (lectotype FI!, designated here).
Cyrtostachys lakka var. singaporensis Becc. (1885: 141).
Type: Singapore,”cultivato nel giardino del Sig. Whampoa”, Anon. (holotype FI!)
Invalid names:
Pinanga purpurea Miq. (1861b: 590), nom. inval., in synon.
Ptychosperma coccinea Teijsm. & Binn. (1866: 69); nom.
nud.
Areca erythrocarpa H. Wendl. in Kerch. (1878: 231),
nom. nud.
Pinanga rubricaulis Linden (1885: 61), nom. nud.
Slender, clustering tree palm with up to c. 3 or more
adult stems up to 15 (– 20) m high. Stem c. 6 – 10 cm
diam., green with greyish stripes or yellow with
somewhat greenish and purplish stripes, internodes
15 – 24 cm long, crown appearing shuttle-cock
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
90
shaped. Leaves 7 – 10 in crown, erect, stiff, to 150 cm
long; sheath tubular, c. 100 cm long, forming distinct
crownshaft, scarlet to bright red, with scattered black
thick scales; petiole elongate, 5 – 50 cm long, 1.5 –
2.5 cm wide and 1 – 2 cm thick at the base, channelled
adaxially, rounded abaxially, red, indumentum as
sheath; leaflets regularly arranged, leathery, 26 – 40
leaflets on each side, 56 – 107 × 3 – 6 cm at middle
portion, apical leaflets 10 – 20 × 1 – 2 cm, briefly
pointed with long tip and sometimes notched at
apices, green, discolorous when dried, glaucous adaxially, waxy white abaxially, mid-vein with discontinuous
membranous brown scales. Inflorescence strongly divaricate, to 90 cm long, branched to 2 (possibly 3) orders,
creamy, green to dark purplish-red; peduncle 5 – 8 cm
long; rachilla 27 – 73.5 cm long and 4 – 6 mm diam.,
calyx persistent on rachillae when fruits fallen off; pits
2 – 5 mm diam., 5 – 7 pits per 1 cm rachilla length.
Staminate flowers 2 – 2.5 × 2 – 3 mm, asymmetrical;
sepals 1.8 – 2 × 2 mm, imbricate, rounded, strongly
keeled; petals 1 – 2 × 1 – 1.8 mm, triangular, brown at
apex and base; stamens 12 – 15; filaments 0.7 – 1 ×
0.2 – 0.3 mm; anthers 1 – 1.5 × 0.5 – 0.8 mm; pollen size,
long axis 36 – 43 µm, short axis 27 – 33 µm, proximal
wall thickness 1.5 – 2 µm, distal wall thickness not
observed, tectum surface microfossulate-rugulate,
trichotomosulcate grains present; pistillode 0.7 – 1 ×
0.2 – 0.5 mm, trifid. Pistillate flowers 4 – 5 × 3 – 4 mm;
sepals 3 – 4 × 2 – 3 mm, imbricate, strongly keeled,
dark brown to black; petals 3 – 3.5 × 2 – 2.5 mm;
gynoecium 3.5 × 1.5 mm (including three recurved
stigma 0.5 – 1 mm); staminodes circular, 0.5 – 1 mm
height. Fruits 7 – 10 × 4 – 7 mm, ellipsoid to ovoid, light
green becoming black when ripe. Seeds 4 – 5 × 3 – 5 ×
3 – 5, ellipsoid to ovoid. (Fig. 3J – M).
DISTRIBUTION. This is the only species found to the west
of Wallace’s Line, occurring in the southern part of
Thailand, Malay Peninsula, Sumatra and Borneo.
SPECIMENS EXAMINED. THAILAND. Narathiwat Province:
Tak Ban, Phru Kok Daam, March 1985, Niyondham 852
(BKF, K!); Tho daeng, c. 30 km SE of Narathiwat, 50 –
100 m a.s.l., Nov. 1990, Barfod & Ueachirakan 41772
(AAU, BKF, K!, PSU); to Daeng, 75 m a.s.l., Oct. 1996,
Barfod et al. 43888 (AAU!, BKF, PSU). N of Sritamerat,
Ta Samet, c. 50 m a.s.l., Jan. 1928, Kerr 14332 (K!).
SUMATRA. Aceh, Asdat 171 (BO!); Asahan, Polak s.n.
(BO!); Bangka Island, Djebus Teysmann s.n. (BO!);
Bengkalis, Selat panjang, 3 m a.s.l., Nov. 1919, Bequin
457 (L!, BO!); near Indrapura, Korthals s.n. (L!, K!; the
type); Riau, Widyatmoko 399 (BO!); Widyatmoko 400
(BO!); South Sumatra, Dransfield JD 1252 (BO!);
Natuna Island, Mogea 2990 (BO!). MALAY PENINSULA.
Selangor: Telok swamp forest, Klang, March 1968,
Dransfield 713 (K!). SINGAPORE.”cultivato nel giardino del
Sig. Whampoa”, Anon. (holotype FI!). BORNEO. SABAH:
Kudat distr., Pulau Balembangan, NE inner side Telok
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
KEW BULLETIN VOL. 64(1)
Lung, 10 m a.s.l., BCS-EFA-LM et. al. SAN 86702 (K!,
KEP, L!, SAR, SING); Sandakan, Jan. 1921, Wood 1111
(A!, PNH, SAN). SARAWAK: Kuching, Nov. 1866,
Beccari PB 2674 (FI!); 1865 – 1869, Beccari PB 3438
(K!); Bintulu, Sept. 1867, Beccari PB 4038 (FI!); 1929,
Clemens 21377 (A!, BO!, K!); Miri distr., Rian road,
50 m a.s.l., April 1959, Saleh 1214 (K!, L!, S, SAR).
BRUNEI DARUSSALAM: Belait, Labi, km 20 Labi road,
burnt over white sand forest, level land, 50 m a.s.l.,
March 1992, Dransfield JD 7279 (K!); Bukit Bakong,
Oct. 1992, Bernstein 278 (K!); Maruntungan, May 1932,
Keith 2491 (K!). CULTIVATED. Indonesia: North Sumatra, Sibolangit Botanic Garden, 500 m a.s.l., Sept. 1927,
Lörzing 12083 (L!); West Java, Bogor Botanic Garden,
origin from Banka Island, loc. V. K. 37, April – May
1936, Furtado SFN 3/1/68 (BO!, K!, L!, SING); loc. V.
G. no. 4., May 1903, Schoute s.n. (L!); Jan. 1906, n.n.
(L!); n.n. (BO!). Malaysia: Penang Botanic Garden,
Sept. 1900, Curtis 3527 (K!). UK: Royal Botanic
Gardens Kew, Palm House, May 1998, 1982-5882
(K!). Seychelles: Victoria, Mali, Dec. 1971, Elizabeth
111 (K!). Singapore: Singapore Botanic Garden, Lawn
K, Sept. 1929, Furtado s.n. (K!, SING). Thailand:
Peninsular Botanic Garden Khao Chong, Trang,
Barfod s.n. (AAU (photo)!).
HABITAT. Cyrtostachys renda grows in lowland peat
swamp forest, especially in coastal areas, but more
rarely occurs in peat swamps in uplands from 0 –
500 m above sea level.
VERNACULAR NAMES. Thailand: kap daeng, mark-dang
(Thai). Malay Peninsula: pinang rajah (Malay). Sumatra: pinang renda or rende’ (Indrapura); pinang rimbou
(Sibolga); pinang lempiauw or pinang lepiaw (Bangka
island). Borneo: malawaring, raring (Brunei). Trade
names: pinang merah, palem merah, (Malay/Indonesia);
sealing wax palm, lipstick palm (English); hsing hsing yeh
tzu (Chinese); rode palm (Dutch).
USES. This palm has limited traditional uses; stems are
used for flooring and leaves for thatch. It is, however, a
highly desirable and widely cultivated ornamental for
tropical regions.
CONSERVATION STATUS. Vulnerable (VU). See Dransfield & Johnson (1991), Kiew (1991), and Mogea
(1991) for conservation status assessment.
NOTES. Cyrtostachys renda differs from all other species
in the bright red crownshaft and leaf sheath, the
lowest number of leaflets (26 – 40 on each side), the
leaflets being waxy white abaxially, the inflorescence
branched mostly to 2 orders (possibly up to 3), the
tectum surface of pollen rugulate, and its preferred
habitat in lowland peat swamp forest in southern
Thailand, Malay Peninsula, Sumatra and Borneo.
In the protologue of Cyrtostachys renda, no material
is cited although a reference is made to Blume’s
account in Rumphia, published some years later, in
which the Korthals specimen is mentioned (Blume
1838, 1843). We formally designate this specimen as
91
A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE)
lectotype. In the case of C. lakka, Beccari (1885) cited
two specimens, Beccari PB 2674 and 4038, both of
which are extant at FI. Harold E. Moore annotated
Beccari PB 2674 (FI) as lectotype in 1956, but to our
knowledge did not publish this lectotypification. We
have formalised this typification here. The transfer of
Cyrtostachys lakka to a synonym of C. renda has already
been made by Whitmore (1982).
The bright green stems and brilliant red to orange
crownshafts make Cyrtostachys renda a highly desirable
and widely cultivated ornamental. Infraspecific taxa
have been described from cultivation and the number
of these could increase in parallel to horticultural
demand. Ellison & Ellison (2001) introduced two
cultivars, C. renda ‘Apple’ and C. renda ‘Orange
Crownshaft’, followed by Waddel (2002) with his C.
renda ‘Theodora Buhler’. Before them, Tucker (1992)
reported, that in Singapore Botanic Garden grew C.
renda ‘Ruby’ and that it was one of the most treasured
specimens of all, and he also discussed a strange
brown crownshafted form of C. renda in Florida. All
the cultivars of C. renda were described based on
different stem and crownshaft colours.
Guinea, Central Province, Boridi, Sept. 1935, Carr
13136 (holotype B†; isotypes K!, L!).
After re-examination of the type of Cyrtostachys compsoclada Burret, no characters diagnostic of Cyrtostachys
were found, while all the characters fit with Heterospathe. There are no pits, and the rachillae are covered
in thick brown indumentum. In the staminate flowers,
petals and filaments are free (connate or united in
Cyrtostachys), and stamens 6 in number (8 – 15 in
Cyrtostachys). Leaf and leaflets are very small for an
adult palm in Cyrtostachys and the flower colour is said
to be purple — colourful inflorescences and flowers
are known in Heterospathe. Moreover, the cited altitude,
5000 feet (1500 m) a.s.l., is higher than usual for
Cyrtostachys. The highest recorded altitude for Cyrtostachys being 900 m a.s.l. for C. barbata, whereas Heterospathe is most frequently found at submontane to
montane elevations.
Cyrtostachys ledermanniana Becc. (1923: 450). See notes
under Cyrtostachys barbata for the discussion.
Excluded and Uncertain Names
Acknowledgements
Many individuals and institutes contributed to the
completion of this work. We would like to thank the
curators and staff of herbaria A, AAU, BH, BO, BRI,
FI, K, KEP, L, LAE, MAN, PNH, SAN and SING for
access to their specimens, databases and for providing
loans in many cases. Dr Hannah Banks is thanked for
assistance with pollen laboratory work and microscopy.
Dr Sander van der Kaars provided fossil pollen
information. Lucy T. Smith prepared the elegant plates.
This research was conducted while CDH did his
Master’s degree in Institut Pertanian Bogor. CDH
would like to thank the Dransfields, the Bakers and
the staff of the Herbarium, Royal Botanic Gardens,
Kew for their support and hospitality during his recent
visits. Special thanks to Marthinus Iwanggin, Piter
Gusbager, Herkilaus Rumakewi, Tobias Paiki, Krisma
Lekitoo, Piter Matani, Maikel Simbiak, Hermus Indou
for help during fieldtrips to Tanah Papua. Financial
support came from BPPS DIKTI Depdiknas, local
government of Provinsi Irian Jaya Barat and local
government of Kabupaten Manokwari, the BAT Biodiversity Partnership to the PONG project of the Royal
Botanic Gardens, Kew, the Pacific Biological Foundation,
the Sibbald Trust, and the Ethnobotanical Conservation
Organisation for South East Asia (ECO-SEA).
Cyrtostachys ceramica (Miq.) H. Wendl. in Kerch. (1878:
242) = Rhopaloblaste ceramica (Miq.) Burret (1928:
288).
Heterospathe compsoclada (Burret) Heatubun comb. nov.
Cyrtostachys compsoclada Burret, Notizb. Bot. Gart. BerlinDahlem 13 (118): 325 (1936). Type: Papua New
Species numbers are given in brackets and in bold
after collection numbers.
Key: (1) C. bakeri, (2) C. barbata, (3) C. elegans, (4) C.
excelsa, (5) C. glauca, (6) C. loriae, (7) C. renda.
Specimens of Uncertain Affinity
Heatubun 679 & 680 (Cyrtostachys sp.).
This specimen was collected from a plant cultivated at
the Kebun Raya Bogor (Bogor Botanic Garden) in the
lawn XIB. XX, close to the nursery and plant
conservation building. In its reddish sheaths this palm
resembles Cyrtostachys renda somewhat but is nevertheless
quite distinct. It is a large, clustering tree palm growing
to 20 m high with a stem of more than 10 cm diam. with
elongate internodes. The crown is hemispherical in
outline and the leaves bear pendulous leaflets, a short
petiole and a greenish-red to yellowish-red sheath and
petiole with a few green stripes. The inflorescence is
large, branched to 3 orders (never 2-branched). Although the clustering habit and red leaf sheath suggest
characters of C. renda, it is far too robust to match
that species. The taxon could well be a result of
hybridisation between two different species (C. elegans
and C. renda) that have been planted together in the
garden (Dransfield 1999). Further studies are required
before a new hybrid species can be formally described.
List of Specimens Examined and Identified
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2009
92
Anon (Cult. Singapore) s.n. (7)
Baker et al. 1110 (6), 1138 (1); Barfod 478 (6), 482 (6);
Barfod s.n. (7); Barfod & Uechirakan 41772 (7); Barfod et
al. 395 (6), 454 (5), 463 (6), 467 (6), 43888 (7); BCSEFA-LM et al. SAN 86702 (7); Beccari PB2674 (7),
PB3438 (7), PB4038 (7); Bernstein 278 (7); Bequin 457
(7); Brass 7162 (6), 7757 (6), 13707 (2), 13807 (6)
Carr 12253 (6); Clemens 1353 (6), 21377 (7); Croft et al.
LAE 68805 (5); Curtis 3527 (7)
Desianto 01 (6); Derbyshire 867 (6); Derbyshire & Hoogland 8020 (6); Dransfield JD713 (7), JD1252 (7),
JD7279 (7)
Elizabeth 111 (7); Essig & Katik LAE 55009 (5); Essig &
Young LAE 74052 (6)
Furtado SFN 3/1/06 (3), 3/1/28 (3), 3/1/68 (7);
Furtado s.n. (7)
Guppy 235 (6); Gusbager 23 (6)
Heatubun et al. 194 (3), 208 (6), 279 (6), 330 (4), 341
(3), 527 (6), 532 (6), 533 (6), 546 (6), 547 (6);
Hoogland & Craven 10114 (6)
Kajewski 2220 (6); Kanehira & Hatusima 12747 (3),
12851 (3); Katik LAE 62219 (5); Keith 2491 (7); Kerr
14332 (7); Kjaer & Magun 512 (5); Korthals s.n. (7);
Kostermans s.n. (6)
Lörzing 12083 (7)
Maturbongs 654 (6); Meijer Dress 501 (6); Mogea 2990
(7); Moore 9272 (5); Morren & Frodin 3189 (6)
Niyondham 852 (7)
Peekel 444 (6); Polak s.n. (7); Pullen 1692 (6), 7305 (6),
8198 (5), 8212 (5), 8231 (5), 8409 (5)
Saleh 1214 (7); Schoode 2248 (6); Schoute s.n. (7)
Takeuchi 8770 (5); Takeuchi et al. 13217 (5); Teysmann s.
n. (7)
van Royen 4734 (6)
Whitmore BSIP 3945 (6), 4210 (6); Widyatmoko 399 (7),
400 (7); Wood 1111 (7)
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