Family: Arecaceae (Palmae)

One of the larger, most natural, and economically important monocotyledonous families
with about 180 genera and 2600 species; typically plants with aerially unbranched,
woody trunks and large compound leaves (palmately or pinnately dissected) in a
terminal crown, and forming extensive pure colonies (Plate 17D). The family is divided
into five subfamilies and much further subdivided (Dransfield et al., 2008). Nypa is a
distinctive rhizomatous mangrove element in the Old World. It forms the monotypic
subfamily Nypoideae, an early diverging taxon, as supported by its unique morphology
and distinctive anatomy (Tomlinson et al., 2011) and extensive early fossil record,
beginning in the early Cretaceous (Dransfield et al., 2008, p. 76). A number of other
palm species may occur as mangrove associates.
The taxa treated here may be distinguished as follows:
1A. Unarmed, rhizomatous palm, massive axis buried in estuarine mud; leaves
pinnately erect to a height of 7 m; fruiting head globose with numerous fibrous
fruits (Plate 18A, B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nypa fruticans
1B. Spiny, trees or Iianes, leaves 1–3 m at most; fruiting heads paniculate. . . . . . . . . . . . 2
2A. Lianes, supported by extended grapnel-like structures; stem diameter 3–4 cm; fruits
with numerous reflexed, regularly overlapping scales. . . . . . . . . Calamus erinaceus
2B. Stems self-supported (Fig. B.9A), clustered, axes to about 10 cm in diameter,
armed with spines that become erected after leaf fall (Fig. B.9B); fruits not
scaly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oncosperma tigillarium
Additional palms that may be encountered, usually as outliers of swamp communities,
include Bactris major Jacq. (tropical America), Phoenix reclinata Jacq. (tropical
Africa), species of Euterpe, Manicaria, and Mauritia (tropical America), and species
of Raphia (Central America and tropical Africa). Such palms may have an appreciable
tolerance of salt water (e.g., Phoenix paludosa Roxb. in Asia). The only other arbores-
cent monocotyledon that might be mentioned in this context is Pandanus candelabrum
as a constituent of the back-mangrove in West Africa.
Nypa Steck 1757 [De Sagu 15; see Moore 1962 Taxon 11: 164–166]
Nipa Thunberg 1782 [Kongl. Vet. Acad. Nya Hanal. 3: 231]
Nypa fruticans (Thunb.) Wurmb. 1781 [Verh. Batav. Genootsch]
Nipa fruticans Thunberg 1782 [loc. cit. 3: 234]
(Figs 5.4, 5.5, 5.6, 10.1; B.6, B.7, B.8; Plates 17, 18)

180
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 Arecaceae (Palmae) 181

A monoecious palm with a thick, subterranean, somewhat fleshy, dichotomously

branched, rhizomatous axis (Fig. 5.6) up to 70 cm in diameter, with oblique raised leaf
scars; axis and leaf bases usually submerged in mud and inundated at high tide (Fig. 5.5),
rarely exposed by erosion (Fig. 5.7). Each terminal shoot supporting a cluster of erect
paripinnate leaves to 7 m long. Leaf with a terete petiole 1–2 m long and a somewhat
bulbous leaf base, closed at the insertion. Leaf base with a pronounced groove accom-
modating the next youngest leaf, but two grooves at the point of dichotomy (Plates 17E,
18C). Leaflets redupicately folded, numerous (30–40), regularly arranged in the grooved
rachis, and each lanceolate, to 70 cm long, with a single prominent adaxial midrib.
Indumentum absent except for frequent scattered abaxial scales, up to 1.5 cm long,
below the midrib. Inflorescence (Uhl, 1972) on a long peduncle, axillary, arising above
the water surface at high tide; bicarinate prophyll (Plate 17F) and initial series of spirally
arranged bracts, sterile and enclosed by the leaf base; distally 7–9 similar bracts, fertile
and subtending lateral branches, the branches themselves branched to three or even four
further orders. Branches usually somewhat adnate to parent axis but without a conspicu-
ous prophyll. Main axis of inflorescence ending in a globose cluster of congested female
flowers, at first enclosed by a series of short sterile bracts; all lateral axes ending in a
club-shaped spike of densely arranged male flowers (Plate 18F). Flowers (Fig. B.6)
dimorphic and sessile, each subtended by an inconspicuous, even filiform bract. Male
flower (Fig. B.6i, j, k) with six short tepals 4–5 mm long, in two whorls, the outer tepals
somewhat widened above; stamens three, united to form a central column, the anthers
dehiscent extrorsely. Pollen spinous (Fig. 9.1). Female flowers (Fig. B.6c–f) with six
tepals much as in the male flower, but exceeded by the three (or four) separate angular
carpels about 1 cm high, the stigmas sessile, funnel-shaped. Carpel unilocular with a
single basal anatropous ovule. Fruiting head developing into an aggregate of fertile and
partly developed but sterile carpels (Plate 18B, G), the individual fruit an angular drupe
with a smooth, dark brown epicarp, mesocarp fibrous, endocarp thick, the seed grooved
adaxially. Endosperm usually with a central hollow; embryo small, basal (Fig. B.7a, b).
Germination incipiently viviparous (Plate 18D, E; Fig. B.7c, d), essentially hypogeal,
but initiated on the fruiting head with the plumule protruding before the fruit is released
(Fig. B.8b, c).

Embryo Morphology

The configuration of the embryo in the germination process is unusual because it is

seemingly the reverse of that in all other palms (Fig. B.8; Plate 18D). The plumule is the
first structure to appear, which implies that it is the root that functions as a haustorium,
not the cotyledon, as in all other palms. The fruit germinates in a horizontal position so
that the first (scale) and subsequent juvenile (foliage) leaves all become erect by rotation
of the leaf base (Fig. B8e, f). To date, I have not succeeded in unraveling this
morphological conundrum.
In virtually all respects, Nypa contrasts remarkably with other palms, reflecting its
highly adaptive morphology and systematic isolation.

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 182 Family

Figure B.6. Nypa fruticans (Arecaceae-Nypoideae) inflorescence and flower. (a) Distal part of
inflorescence at female stage (x1/5; see Plate 18F). (b) Female flower head with lower bracts
removed (x1/2). (c) Single carpels from different aspects (x3/4). (d) Carpel in L.S. (x3/2).
(e) Tepals, one outer, two inner, from a single female flower (x3). (f) Floral diagram of
female flower. (g) Young male spike still enclosed by bract (x1/2). (h) Male spike at anthesis
(x1/2); apex of next higher order spike protruding on left. (i) Floral diagram of male flower.
(j) Male flower just before anthesis (x9). (k) Male flower in L.S. (x9); staminal column
unextended. Note: the perianth has been described as two whorls (Uhl, 1972); this is most
obvious in the female flower (e) and least so in the male (j). Staminal column (k) extends
about two-fold at anthesis, with the stamens standing above the tepals. (Material from
Kuching River, Sarawak.)

Nomenclature, Taxonomy, and Phylogeny

The monogeneric subfamily Nypoideae. It is considered to represent an independent line of

specialization, and because of its numerous distinctive features, has no obvious close
relatives. Relationships with the Pandanaceae have been suggested on the basis of superfi-
cial similarities of the fruiting head, but the basic morphology of the two forms is quite
different. In its vegetative anatomy, it shows all the diagnostic features of the Arecaceae
(Tomlinson et al., 2011); any attempt to segregate it as a distinct family Nypaceae
should be discounted.

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 Arecaceae (Palmae) 183

Figure B.7. Nypa fruticans (Arecaceae-Nypoideae) vivipary. (a)–(d) Attached fruits of increasing
age to show development of viviparous embryo. (d) Corresponds to stage when fruit is released.
(After Tomlinson, 1971.)
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 184 Family

Figure B.8. Nypa fruticans (Arecaceae-Nypoideae) seedlings. (a) Isolated embryo (x3/2),
corresponding approximately to stage of development in Fig. B.7c. (b) Detail of protruding
plumule (x1). (c) Fruit at time of release (x1/2). (d) Isolated embryo (x1/2) from detached, floating
seed. (e) Established seedling (x1/2). (f) Older seedling (x1/8). (After Tomlinson, 1971.)

Distribution and Ecology

Widely distributed from Sri Lanka, the Ganges Delta, Myanmar, and the Malay Penin-
sula, through Indonesia to New Guinea and the Solomon Islands, northward to the
Philippines and Ryukyu Islands, and southward to north Queensland. Nypa has had a
much wider distribution outside its present range because fossil fruits are known from
the Paleocene in Brazil and the Eocene in Europe, Africa, and India. Its pollen (or at

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 Arecaceae (Palmae) 185

least pollen closely similar) is one of the oldest known identifiable angiospermous
pollens referable to a modern species, being recorded in sediments as early as the Upper
Cretaceous.
In its present range, it is a palm of quiet estuaries or shallow lagoons (Fig. 5.5) into
which fresh water may run. It makes extensive pure stands by virtue of its rhizomatous
dichotomizing habit (Fig. 5.6). It seems to favor brackish waters, often forming a wide
border beyond the fringe of adjacent mangroves or swamp forest (Plate 17D). It does
not occur on shores exposed to high wave action and not in hypersaline conditions.
This plant is an unusual mangrove because of its colonial habit. The axis is normally
buried, but when occasionally exposed, resembles a series of overlapping cowpats. The
stem dichotomy is then evident (Fig. 5.7). Isolated stems that are washed out cannot re-
root, but Chai (1982) suggests that large clumps that retain much of their original root
system may become re-established; the only example of vegetative dispersal in
mangroves.
Nypa has been introduced outside its natural range, notably in West Africa where it is
usually regarded as an invasive species despite its economic potential. In Panama,
Duke (1991b) indicates that the small population there is introduced. It sets viable
seedlings and is apparently pollinated by non-stinging bees (Trigona).

Pneumatophores

It is interesting to reflect on the ability of Nypa to provide an oxygen supply to the

submerged rhizome growing within an anoxic environment. This is mediated by an
unusual mechanism involving mature leaves that function in a way comparable to the
modified root system of other mangroves. This has been investigated in detail by
Chomicki et al. (2014) who provide compelling structural and developmental evidence
that the persistent basal part of the leaf axis in Nypa functions in the same manner as a
pneumatophore. The distal part of the leaf abscises at a precise level so that the stump
of the leaf base persists as a vertical projection of uniform height, becoming sealed
distally by tannin-rich tissues. The surface of the leaf base develops lenticels (possibly
modified stomata) as ports of entry to the internal air space system, which is continu-
ous with that of the stem. Measurements suggest that this function can persist for at
least four years. Hence, part of the success of Nypa in the mangal comes from a
unique set of developmental and functional attributes that are modifications of a
standard palm morphology.

Growth

Growth of the axis seems to be continuous, and disinterred stems show no morpho-
logical discontinuities except for the branch forks. The leaves are spirally arranged on
horizontal axes but are brought into an erect position by unequal growth of the leaf base
(Fig. 5.6). The leaf sheath is a closed tube, as in all palms, but the tube is short and

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 186 Family

extends as a short groove into the base of the terete petiole. Branching as described by
Tomlinson (1971) occurs at regular intervals by an equal dichotomy, which is evident in
the leaf immediately behind the fork, and which has a characteristic double groove and
always occupies the same position relative to the fork (Fig. 5.6a; Plates 17E, 18C).
Architecturally, the palm may be referred to Schoute’s model (Hallé et al., 1978), but
with plagiotropic rather than orthotropic axes (cf. Hyphaene). The result is an axis that
spreads by regular forking of monomorphic axes (Fig. 5.6c,d). No vegetative axillary
meristems are produced. Because the leaves are spirally arranged, but must become
vertical at maturity, there is a peculiar growth reorientation as the leaf matures, differing
for each leaf in the phyllotactic spiral.

Reproductive Biology

Clearly, the reproductive mechanisms and genetic consequences of this palm are
unusual among mangroves because of its extensive clonal spread. Most leaves are
capable of subtending an axillary inflorescence, but not all develop completely. Each
inflorescence is protogynous; the stigmas of the carpellate clusters become receptive
as they are exposed by the separation of the enveloping bracts. Receptivity is
indicated by the stickiness of the stigmatic surface. The pollen is released subse-
quently, as the individual male spikes are exposed in turn, elongating beyond the
enveloping bracts (Plate 18F). In each male flower, the staminal column is extended,
carrying the pollen as a mass onto the surface of the spike; flowers are more or less
synchronous in anthesis.
Essig (1973) records two kinds of insects visiting Nypa inflorescences “in significant
numbers.” First, there are small bees of the genus Trigona; second, small flies of the
family Drosophilidae. I can confirm these observations. Apparently, the flies complete
their life cycle in the somewhat fleshy axes of the branches of the male inflorescence,
because these become extensively damaged by larvae that are presumed to be mainly of
the same flies. Essig suggests that the bees are not important pollinators because they
visit male flowers almost exclusively. The flies visit both types of flower and pollen
becomes attached to them. In cultivation, it is considered necessary to hand-pollinate
flowers to achieve successful fruit set; the ease of this and the clonal habit indicate that
plants are self-compatible.
Fruits may fill a single head completely, but isolated fruits commonly occur in a mass
of partly developed but sterile carpels. The mature fruits are dispersed by water; their
detachment is assisted by the plumule, which extends precociously before the seed is
loosened, often penetrating the fibrous tissue of the inflorescence axis. The peduncle
typically becomes pendulous under the weight of the fruiting head, but it is supported
by water at high tide (Plate 18A). Tidal movement aids in the detachment of mature
fruits. The stranded fruit continues to develop producing two or three scale leaves
before the first bladed leaf (eophyll), which is pinnate (Fig. B.8f). The seedling axis is
horizontal from the start.

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 Arecaceae (Palmae) 187

Economic Uses

Nypa is one of the more commercially valuable plants of the mangroves on a sustained
yield basis (Brown and Merrill, 1920). The leaves make a high quality thatch (attap;
Plate 16C); the immature endosperm provides a jelly-like sweetmeat in Malaya; and in
the Philippines, industrial alcohol is extracted by distillation after fermenting the sugar
tapped from the inflorescence, in the manner of toddy (Fong, 1987a, 1989). It is also
suggested as a fuel crop (Fong, 1987b).
Calamus L. [Sp. Pl.: 325]
See Beccari 1908 [Ann. R. Bot. Gard. Calcutta 11 and Appendix 1913]
A genus of over 370 species, mainly dioecious, spiny, climbing palms with whip-like
grapnels that are either extensions of the leaf tip (cirrus) or sterile adnate inflorescences
(flagellum). Leaves pinnate. Fruit covered with numerous reflexed scales. It is exclu-
sively Indo-Malayan except for C. deeratus in West and Central Africa. In a modern
analysis, several segregate genera have been sunk within an expanded Calamus; if
accepted, this would include over 500 species.
Calamus erinaceus (Becc.) Dransfield 1978 [Kew Bull. 32: 484; see also 1979
Malay. For. Rec. 29:130]
A multiple-stemmed rattan forming thickets on the landward edge of mangal or behind
coastal sand bars. Stems climbing to 15 m, 2–3.5 cm in diameter but up to 6 cm wide
including enclosed sheaths. Sheaths orange to yellowish green, very densely armed with
horizontal or oblique whorls of spines. Petiole at insertion on sheath with a stout knee.
Leaf about 4.5 m long with numerous leaflets up to 40 cm by 2 cm, leaf axis extended
into a cirrus about 2 m long. Inflorescences as lateral panicles from axils of uppermost
leaves, with numerous tubular overlapping bracts, ultimate axes (rachillae) with distich-
ous series of flowers, the male much more dense than the female rachillae. Fruit
globular, about 1 cm in diameter, covered with about 12 vertical series of triangular
scales; embryo minute, endosperm homogenous.
The canes of this species have little commercial use. This species is probably the one
referred to as “Daemonorops leptopus” in Watson (1928).
Oncosperma Bl. 1838 [Bull. Sci. Phys. Nat. Neerl. 1: 64]
(Fig. B.9A, B)
A genus within the subtribe Oncospermatinae (Dransfield et al., 2008), occurring from
Sri Lanka widely in southeast Asia.
Oncosperma tigillarium (Jack) Ridl. 1900
[J. R. Asiat. Soc. Straits Branch 33: 173]
O. filamentosa Bl. 1838–1839 [Rumphia 2: 97]
A tall, many-stemmed palm up to 25 m, but the stems are rather slender (scarcely 10 cm
in diameter) with a crown of pinnate leaves up to 4 m long, each leaf with characteristic
drooping leaflets. Stems armed with flat, sharp, black spines. Both Watson (1928)

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 188 Family

a b

Figure B.9. Oncosperma tigillarium (Arecaceae-Arecoideae). (a) A characteristic back-mangal

species in Bako National Park, Malaysia. (b) Spiny trunk; stems are used in construction after
the spines are shaved off.

and Chai (1982) record this palm as marking the inner limit of the mangrove that
is found in regions transitional to non-littoral forest. Watson indicates that it grows
inland and suggests it is likely to be killed by salt water. It is successfully cultivated at
the Montgomery Botanical Center in South Florida adjacent to tidally influenced
open water.

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