บทความวิจัย (Research Article)
*Corresponding author: [email protected]

Comparative Anatomy of Two Mangrove Species,
Xylocarpus granatum and Xylocarpus moluccensis
(Meliaceae)

Boonyawee Chorchuhirun*, Ekaphan Kraichak and Prasart Kermanee
Department of Botany, Faculty of Science, Kasetsart University, Bangkhen Campus,
Ladyao, Chatuchak, Bangkok, 10900

Received: January 21, 2020; Accepted: February 13, 2020


Abstract
Xylocarpus granatum and Xylocarpus moluccensis are true mangrove species, which are found
in the central zone of mangrove forests. Anatomical character plays important roles to plant functions
and adaptations to cope with certain environments. The plant samples were collected from the Sirinart
Rajini Ecosystem Learning Center, Prachuabkirikhan province. Permanent slides of the samples were
prepared using a paraffin method. The anatomical characters of root, stem, petiole, petiolule, rachis
and leaf blade were examined and compared. The results showed that X. granatum and X. moluccensis
shared their anatomical characters, i.e., tanniferous cells and crystals are common in both species.
However, there are some different characteristics such as primary xylem in root, shape of vascular
bundle in petiole, petiolule and leaf blade including number of hypodermis and palisade layers. Overall
of this study, the anatomical character can discriminate between the two species.

Keywords: anatomy; mangrove; Xylocarpus granatum; Xylocarpus moluccensis

1. Introduction
Mangroves are evergreen forests, which
are found along coastlines, shallow-water
lagoons, estuaries or rivers in tropical and
subtropical areas. The mangrove ecosystem
represents interphase between terrestrial and
marine communities, which receive strong
winds, sea tides and water salinity (FAO, 2007;
Giesen et al., 2007). Mangrove plants have
several mechanisms of adaptation to an
anaerobic condition in saline water, including
rooting system as morphological adaptation,
pneumatophores as anatomical adaptation and
water conservation as physiological adaptation
(Göltenboth et al., 2006).
The variation of salinity, elevation, wave
exposure and tidal inundation lead to the
zonation of mangroves. The mangrove zonation
is classified into 4 main zones. The seaward
zone is inundated by all high tides. A mid zone DOI: 10.14456/tjst.2020.25

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is inundated by medium-high tide. The landward
zone is inundated by the highest tides only.
Brackish stream mangroves are found along
brackish to freshwater streams (Giesen et al.,
2007; Waycott et al., 2011). The landward zone
is interesting for a study about mangrove plant
adaptations because it is a transitioning
ecosystem between mangrove and terrestrial
communities. The outstanding true mangrove
plants at the landward zone is Xylocarpus. This
genus is important for traditional medicine.
Which used fruits and seeds of Xylocarpus
granatum J. Koenig and X. moluccensis (Lam.)
M. Roem for treating diarrhea and bark
decoction for treating cholera. Moreover, X.
granatum woods are attractive and used for
furniture making (Haron and Taha, 2010; Das et
al., 2014; Baba et al., 2016).
X. granatum is a medium sized evergreen
tree. Compound leaves have 1-2 pairs of
leaflets. Their leaflets are rounded and
coriaceous. The trunk is smooth with light brown
outer bark, which peels off with thin flakes, and
red inner bark. Buttress roots, which are ribbon-
like plank forms, are surrounded the trunk.
Florets include 4 white petals and 4 yellowish-
green sepals. Fruits are yellowish- brown
globose up to 25 cm diameter (Haron and Taha,
2007; Haron and Taha, 2010; Das et al., 2014;
Ragavan et al., 2014; Ragavan et al., 2015).
X. moluccensis is a medium sized
deciduous tree. Compound leaves, which
arranged spiral or opposite, have 1-3 pairs of
leaflets. The trunk is rough with dark brown outer
bark, which peels in long narrow strips.
Pneumatophores are conical shape up to 30 cm
tall. Inflorescence and florets are similar to X.
granatum. Fruits are brownish green sub-
globose up to 12 cm diameter. Both species
have buoyant fruits and seeds (Haron and Taha,
2010; Das et al., 2014; Ragavan et al., 2014;
Ragavan et al., 2015).
Morphology and anatomy play important
roles in adaptation to the stress environment in
the mangrove forest, but there are a few pieces
of knowledge about them (Das and Ghose,
1996; Haron and Taha, 2007; Rodriguez et al.,
2012). This research focused on anatomical
characters of root, stem, petiole, rachis, petiolule
and leaf blade in the genus Xylocarpus found in
Thailand. The obtained data may support
species identification and to understand the
anatomical and physiological adaptations of
Xylocarpus.

2. Materials and Methods
The specimens of X. granatum and X.
moluccensis were collected from a mangrove
forest at the Sirinart Rajini Ecosystem Learning
Center in Pranburi, Prachuabkirikhan province.
The six replicates of roots at 1 cm from the root
tip, stem at 2-4 cm from shoot tip and mature
leaves at 5
th
node from the apex were fixed in
formalin acetic alcohol (FAA) solution.
Permanent slides were prepared using the
paraffin method (Johansen, 1940; Kermanee,
2008) . The samples were dehydrated with
tertiary butyl alcohol series, infiltrated in liquid
paraffin, embedded in paraplast and sectioned
into 10-15 microns thickness. The sections were

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stained with safranin-T and fast green
combination and then were mounted with
permount. The specimens were examined with
a compound light microscope (Zeiss; Axioskop
2, Germany) assembled with a camera (Zeiss;
AxioCam MRc, Germany) and Zen 2 program.

3. Results
Six organs of X. granatum and X.
moluccensis including root, stem, petiole, rachis,
petiolule and leaf blade were examined. The
anatomical characters were described as
follows:
3.1 X. granatum
3.1.1 Root
The outer layer is epidermis which
accumulates suberin (Figure 1A). There is a
layer of exodermis beneath the epidermis.
Cortex composes of parenchyma with air
spaces. Tanniferous cells were observed in the
cortex. A layer of endodermis with Casparian
strip surrounds a stele. The stele is actinostele
with 7 arches of primary xylem. The center of
the root is pith with some tanniferous cells
(Figure 1B).
3.1.2 Stem
The outer layer is periderm,
consisting of several layers of phellem. The
cortex contains parenchyma. A number of
tanniniferous cells and druse crystals were found
in the cortex. Stele presents secondary phloem
and secondary xylem. Gelatinous fibers were
observed in phloem and xylem (Figure 1C and
1D).
3.1.3 Petiole
The outer layer is periderm which
consists of phellem, phellogen and phelloderm.
Cortex composes of parenchyma with
tanniferous cells and druse crystals (Figure 1E).
The vascular bundles arranged in a semi-circular
shape.
3.1.4 Rachis
The outer layer is the epidermis.
Some epidermal cells deposit suberin. There is
a layer of periderm beneath the epidermis. The
cortex contains parenchyma which some cells
contain tannin. Stele presents a ring of
secondary phloem and secondary xylem (Figure
1F).
3.1.5 Petiolule
The outer layer is epidermis which
accumulates suberin. Periderm presents
beneath the epidermis. The cortex consists of
parenchyma which stores druse crystals. The
vascular tissues are collateral bundles which
arrange in semi-circular shape (Figure 2A and
2B).
3.1.6 Leaf blade
Midrib composes of the upper and
lower epidermis which are covered with thick
cuticle (2.66±0.29 µm). The cortex consists of
round-elliptic parenchyma. Abundant druse
crystals are found in the cortex. Vascular
bundles contain groups of collateral bundles in
semi-circular shape (Figure 2C and 2D). The
mature blade is bifacial, composes of upper and
lower epidermis. There are 2-3 layers of
hypodermis under the epidermis (Figure 2E).
Typical stomata are observed only on the lower
epidermis (Figure 2F). Mesophyll composes of 2

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types of modified parenchyma. Palisade
mesophyll appears on the upper part of leaf
blades, consisting of 2-3 layers. Spongy
mesophyll presents in the lower part with large
intercellular spaces. Druse crystals are found
in







Figure 1 Transverse sections showing anatomical characters of X. granatum; (A) Root (B) Stele of
root (C) Stem (D) Periderm, cortex, phloem, xylem and pith of stem (E) Petiole (F) Rachis
(C. = cortex, D.Cr. = druse crystal, E. = epidermis, En. = endodermis, Ex. = exodermis,
G.F. = gelatinous fiber, Pe. = periderm, Ph. = phloem, Pi. = pith, Xy. = xylem)

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Figure 2 Transverse sections showing leaf blades anatomy of X. granatum; (A) Petiolule (B) Druse
crystal in petiolule (C) Leaf blades (D) Midrib (E) Blade (F) Lower epidermis with typical
stomata (C. = cortex, D.Cr. = druse crystal, E. = epidermis, H. = hypodermis, L.E. = lower
epidermis, P.M. = palisade mesophyll, Pe. = periderm, Ph. = phloem, Pi. = pith, S.M. = spongy
mesophyll, T.S. = typical stoma, U.E. = upper epidermis, V.B. = vascular bundle, Xy. = xylem)

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Figure 3 Transverse sections showing anatomical characters of X. moluccensis; (A) Root (B) Stele of
root (C) Stem (D) Epidermis, cortex, phloem and xylem of stem (E) Petiole (F) Rachis
(C. = cortex, D.Cr. = druse crystal, E. = epidermis, En. = endodermis, Ex. = exodermis,
G.F. = gelatinous fiber, Pe. = periderm, Ph. = phloem, Pi. = pith, Xy. = xylem)

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Figure 4 Transverse sections showing leaf blades anatomy of X. moluccensis; (A) Petiolule (B) Druse
crystal in petiolule (C) Leaf blades (D) Midrib (E) Blade (F) Lower epidermis with typical
stomata (C. = cortex, E. = epidermis, H. = hypodermis, L.E. = lower epidermis, P.M. = palisade
mesophyll, Pe. = periderm, Ph. = phloem, Pi. = pith, S.M. = spongy mesophyll, T.S. = typical
stoma, U.E. = upper epidermis, V.B. = vascular bundle, Xy. = xylem)

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spongy mesophyll.
3.2 X. moluccensis
3.2.1 Root
The outer layer is epidermis which
accumulates tannin (Figure 3A). The stele is
actinostele with 4 arches of primary xylem. The
center of the root is pith (Figure 3B). Exodermis
beneath epidermis, cortex and endodermis with
Casparian strip are similar to X. granatum.
3.2.2 Stem
The outer layer is epidermis which
accumulates suberin. The cortex consists of two
sizes of parenchyma. Tanniferous cells were
observed in the cortex. Stele and gelatinous
fibers present similar to X. granatum. (Figure 3C
and 3D).
3.2.3 Petiole
The outer layer is periderm which
consists of phellem, phellogen and phelloderm.
Cortex composes of parenchyma with
tanniferous cells and druse crystals. The
vascular bundles arrange in cordate shape
(Figure 3E).
3.2.4 Rachis
Suberin accumulation is observed
in the epidermal layer. The cortex consists of two
sizes of parenchyma. Tanniferous cells are
observed. The stele is similar to X. granatum
(Figure 3F).
3.2.5 Petiolule
The outer layer is periderm. Druse
crystals are observed in the cortex. The vascular
tissues are collateral bundles, which arrange in
cordate shape (Figure 4A and 4B).
3.2.6 Leaf blade
Midrib cortex consists of round-
elliptic parenchyma with druse crystals and
tanniferous cells. Vascular bundles contain
groups of collateral bundles arranging in oval
shape (Figure 4C and 4D). The mature blade is
bifacial, composes of the upper and lower
epidermis with thick cuticle (3.22±0.49 µm).
There is a layer of hypodermis under the
epidermis (Figure 4E) . Typical stomata are
observed only on the lower epidermis (Figure
4F) . Mesophyll composes long palisade
mesophyll (84.67±7.82 µm) appears on the
upper part of leaf blades, consisting of 1-2
layers. Spongy mesophyll presents large
intercellular spaces which cells deposit druse
crystals.

4. Discussion
4.1 Inclusion in Xylocarpus
Tanniferous cells are common in the
genus Xylocarpus. Tannin is a derivative of
phenolics (Alongi, 1987; Amarowicz, 2007)
which plays an important role in preventing
bacterial and fungal infestations (Giner-chavez,
1996) and alleviates damage in tissue caused
by ion excess, salt or hydrogen sulfide toxicities
(Kimura and Wada, 1989). Interestingly, the
distribution of tanniferous cells in the two species
is different. For instance, they were found in the
root cortex of X. granatum but absent in X.
moluccensis (Table 1). However, they were
observed in the leaf tissue of X. moluccensis.
This implied that the leaf of X. moluccensis is
more susceptible to salt than root tissue.
Crystals are observed in most parts of

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the species. Crystals are calcium oxalate
compound which is derived from metabolic
waste (Arnott and Pautard, 1970, Franceschi
and Hornor, 1980). Their roles as making the
plants inedible to predators and detoxification
(Rasmussen and Smith, 1961; Finley, 1999).
4.2 Root
The primary roots of both species
showed different characters. The primary xylem
of X. moluccensis is tetrarch (4 arches of xylem)
while X. granatum is polyarch (7 arches of
xylem) . This character can be used to
discriminate between the two species. Roots of
both species contain pith tissue in the stele. This
is not found in normal eudicots. However, there
are reports that found this character in other
mangrove species, including Rhizophora mangle
(Menezes, 2006), R. apiculata, R. mucronata
(Thonglim, 2018) and Bruguiera gymnorrhiza
(Pornpromsirikul, 2017) . The character was
mentioned to aerial roots by Gill and Tomlinson
(1977), these roots had an origin at the trunk or
the other aerial roots (adventitious roots). This
contrasts with X. granatum that its roots
germinated from radicle (Das and Ghose, 2003).
Polyarch xylem presented high vessel grouping
and high vessel density, which were adapted to
reduce the risk of air bubble formation in xylem
sap (Baas et al., 1983; Cochard, 2006; Robert
et al., 2009).
4.3 Stem
The epidermal layer of X. moluccensis
stem deposited suberin while X. granatum
developed a periderm layer. The major
component of periderm is phellem tissue, which
contains suberin. Suberin is an insulating
substance that can exclude salt from the
environment to enter the plant body. The
important function of periderm is the plants’
tissue protection (Esau, 1965; 1977). Abundant
gelatinous fibers were observed in the phloem
(phloem fibers) and xylem (xylem fibers) .
Gelatinous fiber is a labriform fiber having
protoplasm and lacks lignin on its cell wall. It
possesses a flexible property. As Xylocarpus
grows under windy areas, the gelatinous fibers
can prevent stem breaking (Tomlinson, 2003;
Pramod et al., 2014).
4.4 Petiole and petiolule
Petiole and petiolule of both species
present periderm, secondary phloem and
secondary xylem. This means that they possess
secondary growth like a stem. Vascular bundles
of X. granatum arrange in semi-circular shape
while the cordate shape is observed in X.
moluccensis. This character would be support
species identification.
4.5 Leaf blade
Xylocarpus leaves are covered with a
thick cuticle. The cuticular layer can provide a
hydrophobic barrier to water loss (Rotond et al.,
2003; Reef and Lovelock, 2015) . Cortex at
midrib of X. granatum composes of normal
parenchyma, while X. moluccensis consists of
parenchyma with tanniferous cells. Druse
crystals were found in the midrib of both species.
Vascular bundles of X. granatum is a semi-
circular shape, while X. moluccensis is oval
shape. Beneath the upper epidermis is a
hypodermis, consisting of 2-3 layers in X.

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granatum, whereas X. moluccensis has only one
layer. Xylocarpus grows under several stresses
including saline water, strong wind and too much
sunlight. The hypodermis is a xerophytic
character. Its functions are water storage and
infiltrate the excess light intensity (Saenger,
2002). Typical stomata are observed only on
lower epidermis of both species which differs
from a previous study of Haron and Taha (2010)
that reported sunken stomata in Xylocarpus sp.
This character responses to reducing water
evaporation in a rigid environment (Saenger,
2002). Palisade mesophyll consisting of 1-3
layers of chlorenchyma with less intercellular
spaces. The palisade cells in X. moluccensis is
longer (84.67 ± 7.82 µm) than in X. granatum
(23.73 ± 2.80 µm). The long palisade cells of
mangrove plants help to increase
photosynthesis efficiency (Thonglim, 2018).
The comparison of distinct anatomical
characters in X. granatum and X. moluccensis is
summarized in Table 1.

Table 1 Comparison of distinct anatomical characters in Xylocarpus granatum and Xylocarpus
moluccensis

Organs
Mangrove species
X. granatum X. moluccensis
Root
Primary xylem with 7 arches Primary xylem with 4 arches
Tanniferous cells in cortex No tanniferous cells in cortex
Stem One zone of cortex Two zones of cortex
Petiole
Semi-circular shape of vascular bundles Cordate shape of vascular bundles
Suberin accumulation in epidermis layer Present periderm
Petiolule Semi-circular shape of vascular bundles Cordate shape of vascular bundles
Rachis Present periderm Absent periderm
Leaf
Semi-circular shape of vascular bundles in midrib Oval shape of vascular bundles in midrib
No tanniferous cells in cortex Tanniferous cells in cortex
2-3 layers of hypodermis 1 layer of hypodermis
2-3 layers of palisade cells 1-2 layers of palisade cells

5. Conclusion
5.1 X. granatum and X. moluccensis
share anatomical characters but there are some
different characteristics such as primary xylem in
the root, shape of vascular bundle in the petiole,
petiolule and blade, number of hypodermis and
palisade layers. These can be used to
discriminate the two studied species.
5.2 Tanniferous cells and druse crystals
are common in all parts of both species.

6. Acknowledgements
The authors wish to acknowledge the
Sirinart Rajini Ecosystem Learning Center for co-
operating in specimen collection. The research
was financially supported by the Development

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and Promotion of Science and Technology
Talents Project (DPST).

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