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FIRENZE
UNIVERSITY
PRESS
June 2020
Vol. 75 - n. 1
0083-7792
WEBBIA
Journal of Plant
Taxonomy and Geography
Poste Italiane spa - Tassa pagata - Piego di libro
Aut. n. 072/DCB/FI1/VF del 31.03.2005
Journal of Plant Taxonomy
and Geography
WEBBIA
Vol. 75, n. 1 - 2020
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Webbia. Journal of Plant Taxonomy and Geography
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Cover image: Nepenthes latiana M.N. Faizal, A.Amin & N. Dome (see in this issue: Mohd Norfaizal Ghazalli et al.)
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Webbia. Journal of Plant Taxonomy and Geography 75(1): 5-28, 2020
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WEBBIA
Journal of Plant
Taxonomy
and Geography
ISSN 0083-7792 (print) | ISSN 2169-4060 (online) | DOI: 10.36253/jopt-7950
Citation: Mohd N. Ghazalli et al.
(2020) Nepenthes latiffiana and N.
domei (Nepenthaceae), two new spe-
cies of pitcher plants from Terengganu,
Peninsular Malaysia. Webbia. Journal
of Plant Taxonomy and Geography
75(1): 5-28. doi: 10.36253/jopt-7950
Received: January 29, 2020
Accepted: April 30, 2020
Published: June 30, 2020
Copyright: © 2020 Mohd Norfaizal
Ghazalli, Amin Asyraf Tamizi, Dome
Nikong, Edward Entalai Besi, Muham-
ad Ikhwanuddin Mat Esa, Anuar Rasy-
idi Mohd Nordin, A. Latiff, Ahmad Zaki
Zaini, Mohamad Alias Shakri. This is
an open access, peer-reviewed arti-
cle published by Firenze University
Press (http://www.fupress.com/webbia)
and distributed under the terms of the
Creative Commons Attribution License,
which permits unrestricted use, distri-
bution, and reproduction in any medi-
um, provided the original author and
source are credited.
Data Availability Statement: All rel-
evant data are within the paper and its
Supporting Information les.
Competing Interests: The Author(s)
declare(s) no conict of interest.
Editor: Riccardo M. Baldini, University
of Florence, Italy
Nepenthes latiana and N. domei
(Nepenthaceae), two new species of pitcher
plants from Terengganu, Peninsular Malaysia
M N G,*, A A T, D N,
E E B, M I M E, A R-
M N, A. L, A Z Z, M A
S
1 Resource Utilisation and Agrobiodiversity Conservation Programme (BE2), Agrobiodi-
versity and Environment Research Centre, Malaysian Agricultural Research and Develop-
ment Institute (MARDI) Headquarters, 43400 Serdang, Selangor, Malaysia
2 Agri-Omics and Bioinformatics Programme (BN1), Biotechnology and Nanotechnology
Research Centre, Malaysian Agricultural Research and Development Institute (MARDI)
Headquarters, 43400 Serdang, Selangor, Malaysia
3 DigitalDome Photography, 21500 Permaisuri, Terengganu, Malaysia
4 Department of Biology, Faculty of Science, Universiti Putra Malaysia (UPM), 43400
Serdang, Selangor, Malaysia
5 School of Environmental Science and Natural Resources, Faculty of Science and Tech-
nology, Universiti Kebangsaan Malaysia (UKM), 43600 Bangi, Selangor, Malaysia
6 Electron Microscopy Unit, Faculty of Science and Technology, Universiti Kebangsaan
Malaysia (UKM), 43600 Bangi, Selangor, Malaysia
7 Fox Extreme Photography, 21200 Kuala Terengganu, Terengganu, Malaysia
*Corresponding author. Email: mnfaizal@mardi.gov.my
Abstract. Two new species of Nepenthes from Terengganu, Peninsular Malaysia, N. latif-
ana M. N. Faizal, A. Amin & N. Dome and N. domei M. N. Faizal, A. Amin & A. Lat-
i, are described and illustrated.
Keywords: pitcher plants, carnivorous plants, Nepenthes, Nepenthes domei, Nepenthes
latiana, Terengganu, Peninsular Malaysia.
INTRODUCTION
In Peninsular Malaysia, a total of eleven species of pitcher plants (Nepen-
thes) or locally known as “periuk kera” have been recorded (Jebb and Cheek
1997; Cheek and Jebb 2001; Clarke 2001; McPherson 2009). e rst taxo-
nomic account of the genus in Peninsular Malaysia was given by Ridley
(1924) who recorded a total of ten species, namely N. ampullaria Jack., N.
albomarginata Lobb., N. gracillima Ridl., N. alba Ridl., N. ramispina Ridl.,
N. sanguinea Lindl., N. macfarlanei Hemsl., N. gracilis Korth., N. raesiana
Jack and N. phyllamphora Willd. (= N. mirabilis (Lour.) Druce). Aer Ridley’s
6Mohd Norfaizal Ghazalli et al.
treatment, there have been some further researches on
the genus by Holttum (1940), Kiew (1990), Turner (1995),
Clarke (2002), Adam et al. (2005) Lati et al. (2011),
Clarke and Lee (2012) and Lati and Norsiah (2016)
regarding the diversity, distribution, and ecological
study of Nepenthes species and natural hybrids. Another
taxonomic study of the Peninsular Malaysian taxa was
carried out by Rohana in 1988 that covers basic ana-
tomical features and systematic study of the species and
hybrids. Turner (1995) listed ten species, which instead
included two natural hybrids, namely N. × hookeri-
ana and N. × trichocarpa, with N. alba and N. ramispi-
na being excluded. Clarke (1999) added N. benstonei, a
species then known from Bukit Bakar, Kelantan, to the
list and the latest was Adam and Hamid (2007) who
described N. sharifah-hafsahii which happens to be a
natural hybrid between N. gracilis and N. mirabilis rath-
er than a true species (McPherson 2009).
e most comprehensive and well accepted taxo-
nomic enumeration works of the genus were given by
Jebb and Cheek (1997), Cheek and Jebb (2001) and
Clarke (2001) who recognised eleven species. In the
course of continuous eld trips to the state of Tereng-
ganu, the authors had encountered several populations
of pitcher plants in Setiu whose features did not match
the currently described taxa of Peninsular Malaysia.
From this nding, we further analysed the morphologi-
cal, anatomical and micromorphological characteris-
tics and have discovered two distinct new species (as
described in this article). e descriptions of N. domei
and N. latiana herein can be considered as a compre-
hensive work in the taxa identication as the sheet con-
tains the sterile/fertile parts and pitchers, anatomical
and micromorphological evidences, and enumeration
of prominent plant characteristics. We also conducted a
comparative molecular study on N. domei using internal
transcribed spacer (ITS) DNA region to further dieren-
tiate it from a closely related species.
MATERIALS AND METHODS
Analyses of morphological characteristics on
new Nepenthes specimens were based on living plants
observed in situ and also from herbarium type speci-
mens (MDI12424 and MDI12423) deposited at MARDI
Herbarium (MyGenebank™ Complex MARDI Serdang).
For leaf anatomy analysis, collected and described
Nepenthes leaves were xed in AA solution (1 Acetic acid
: 3 Alcohol). Investigated leaf parts including midrib,
lamina and margin were sectioned in a range of thick-
ness from 15-30 m using sliding microtome. e leaves
then were cleared using bleaching agent, washed in vari-
ous alcohol series and stained in Safranin and Alcian
blue for 5 minutes each before proceeded for dehydra-
tion stage through an ascending alcohol series (50-
100%). During the nal dehydration stage, the leaf speci-
mens were dierentiated in 70% alcohol with a drop of
hydrochloric acid (HCL) before mounted in Euparal.
Specimen slides were kept in the oven for two weeks at
about 60-70°C. Photographs of the investigated sections
were taken using Olympus SZH40 microscope and the
images were processed using CellSens Image Analysis
soware.
For micromorphology study, Nepenthes epidermis
preparative fragments about 3 mm long were excised
from the middle portion of approximately three mature
leaves of each species. Cuticle parts were prepared by
soaking leaf blade fragments in 5-10% aqueous chromi-
um trioxide until all organic material except the cuticle
was dissolved. Targeted leaf cuticles were mounted on
aluminium stubs with double-sided adhesive tape and
proceeded for air drying. e stubs were then sputter-
coated with pure gold to a maximum thickness of 15
nm and examined with LEO (Model 1450 SEM) - Field-
Emission Scanning Electron Microscope (FESEM). e
remaining cuticles were soaked in 5% ammonia and
mounted on microscope slides in Canada balsam.
Results of the adaxial and abaxial epidermal lay-
ers under scanning electron microscope (SEM) and leaf
anatomical study of N. domei, N. latiana, N. benstonei
and N. sanguinea are shown in Figure 5 to 12 and sum-
marised in Table 2 and 3. ese results are combined
with transverse section of the leaf lamina and margin
showing epidermal cells, mucilaginous idioblast, glan-
dular and simple trichomes, solitary crystals observation
on the parenchyma and along a vein, and signicant
venation characters. e systematic signicant of leaf
anatomy and micromorphology evidences dierentiat-
ing these investigated Nepenthes species are presented in
Table 2 and 3.
Finally, for molecular study, all genomic DNA puri-
cations were done according to DNeasy Plant Mini Kit
(Qiagen) manufacturer’s protocol and matured leaves
(100 mg per sample) from N. domei sp. nov. (Tereng-
ganu), N. benstonei (Bukit Bakar, Kelantan) and N. san-
guinea (Fraser’s Hill, Pahang) were used. e internal
transcribed spacer (ITS) partial gene (~610bp) was PCR-
amplied from Nepenthes DNA using universal ITS1-2
primers (ITS_F: 5’-AGGAGAAGTCGTAACA AGGTT;
ITS_R: 5’ GATGCAACCTTGGCCTT) and Q5 High-
Fidelity DNA polymerase (New England Biolabs, UK).
e parameters for thermocycler were set as follows: ini-
tial denaturation at 98°C for 30s, denaturation at 98°C
7
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
for 10s, annealing at 61°C for 15 s, extension at 72°C for
20 s, and nal extension at 72°C for 2 min. e ampli-
cons were sent for Sanger sequencing and the sequencing
data was trimmed and aligned using BioEdit Sequence
Alignment Editor. Phylogenetic inference of nine pen-
insular Nepenthes species (including N. domei sp. nov.)
was reconstructed in Molecular Evolutionary Genet-
ics Analysis soware (MEGA5) using Neighbor-Joining
(NJ) method (1000 bootstraps) based on the ITS par-
tial sequences isolated in this study as well as sequences
obtained from the NCBI database (Bunawan et al. 2017;
Alamsyah and Ito 2013; Renner and Specht 2011).
TAXONOMIC TREATMENT
Nepenthes latiana M. N. Faizal, A. Amin & N. Dome,
sp. nov. (Figures 1 & 2)
Type: Malaysia, Peninsular Malaysia, Terengganu, Setiu,
30 April 2019, MNFG751 (MDI!), MDI12424, Mohd. Nor-
faizal, Amin Asyraf, Dome Nikong, Muhamad Ikhwan,
Edward Entalai & Anuar Rasyidi, [holotypus, MDI]
Diagnosis
Nepenthes latiana diers from N. sanguinea in peri-
stome morphology which is considerably developed,
loosely cylindrical, with expanded outer margin part
towards both sides of the mouth forming ap-like struc-
ture which is especially prominent in upper pitcher (vs.
simple, expanded, outer margin usually markedly sinu-
ate where the peristome is widest); climbing stem with
simple hairs, cross section cylindrical (vs. glabrous,
sharply angular) and lid margin slight wavy to wavy that
retains its morphology in dried and wet preserved speci-
mens (vs. at).
Description
Terrestrial climber to 0.5-1.5 m tall. Climbing stems
cylindrical to rounded, particularly towards the leaf
nodes, 0.8-1.2 cm in diameter. Internodes 2.2-3.5 cm
long. Leaves coriaceous, sessile to less amplexicaul, lin-
ear-lanceolate, apex obtuse to emarginated, 15-20 cm
long, 5-6 cm wide; base clasping stem for about ½ of
its circumference; longitudinal veins 1-3 on each side
of the midrib, pennate nerves conspicuous; tendril with
slender formation in the middle, 10-14 cm long, with
minute hairs. Lower pitchers up to 18-26.5 cm long, 4.9-
8 cm wide, eshy coriaceous texture, arising abruptly
from the tendril; broad cylindrical at the lower part of
the pitcher, ½ upper of the pitcher slightly swollen with
hip formation, 9 cm length, up to 7 cm above of the hip
till nearly closer to the mouth; cylindrical above, with
slight widening towards the mouth, surface of the hip
smooth; two fringed wings up to 0.9-1.0 cm wide, runs
along the pitcher and widest at the mouth, with multi-
cellular hair elements up to 0.3-0.6 cm long; spur with
branched form, reaching 1.0 cm long; mouth rounded;
peristome loosely cylindrical but widening or expanded
towards lateral sides of the pitcher mouth (ca. 1.5 cm
wide each side), outer surface smooth, reaching 1.1 cm
wide (in front of the mouth), ribs (at the part of the peri-
stome) reaching 1.6 cm wide; lid rounded, up to 6.5 cm
long and 6 cm wide, base retuse, without appendages or
trichomes on the lower surface of the lid, ~105-126 dot-
ted nectar glands superuous and dense at the center
and base of the lid. Upper pitchers up to 20-23.5 cm long,
5-7.5 cm wide, coriaceous texture, arising slight abruptly
from the tendril; broad cylindrical at the ¼ lower part
of the pitcher, ¾ upper of the pitcher cylindrical and
slight narrowing towards the mouth; with minute wings;
mouth ovoid to sub ovoid, peristome cylindrical but
with expanded lateral sides (1.3-1.4 cm wide each side)
and attened towards the front of the mouth part, outer
surface smooth, reaching 9mm wide, ribs reaching 1.3
mm wide; lid rounded, up to 4.8-5.2 cm long and 5.5 cm
wide, base cordate, without appendages or trichomes on
the lower surface of the lid. Male inorescence a raceme,
peduncle up to 16 cm, rachis up to 40 cm, partial pedun-
cles single-owered, bracts absent, pedicel 10-13 mm
long, tepal ovate, up to 2-5 mm, staminal column 2.5-
6 mm long, anther head 2-3.75 mm. Colour of the living
plants: Lower pitcher combination of green-yellowish-
brown hues with several red blotches formation on the
1/3 of the upper part of the pitcher body. Upper pitcher
light green with fade specks. Pitcher interior green-
yellowish to reddish. Leaves green. Stem green to dark
green. Colour of the dried specimen light brown – pale
brown (lamina, stem and pitcher).
Etymology
Named in honour of Emeritus Professor Dato’ Dr. A.
Lati Mohamad, a prominent gure in Malaysian Bot-
any Field from Universiti Kebangsaan Malaysia, Bangi
Campus, Malaysia.
Distribution and habitat
Nepenthes latiana is currently known only from the
restricted area at the type locality in Setiu, Terengganu.
Ecology
Nepenthes latiana occurs as terrestrial climbers on
hilly terrains with slight shaded, upper hill forest habi-
tats, at an altitude of c. 1000-1100 m.
8Mohd Norfaizal Ghazalli et al.
Figure 1. Nepenthes latiana, in situ: (A) Lower pitchers. (B) Upper pitcher. (C) Immature upper pitchers. (D) Lid, peristome and waxy
inner surface of lower pitcher. (E) Habit showing the lower and upper pitchers. Photographs by Mohd Norfaizal Ghazalli and Amin Asyraf
Tamizi.
9
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 2. Line drawing of Nepenthes latiana: (A) Habit, with inorescence. (B) Leaf blade. (C) Upper pitcher, viewed from side with
spur. (D) Upper pitcher frontal view. (E) Lower pitcher. (F) Adaxial and abaxial side of the lid. (G) Male ower. Based on 12424 (MDI). All
drawn by Khalid Hashim.
10 Mohd Norfaizal Ghazalli et al.
Nepenthes domei M. N. Faizal, A. Amin, & A. Lati, sp.
nov. (Figures 3 & 4)
Type: Malaysia, Peninsular Malaysia, Terengganu, Setiu,
30 April 2019, MNFG750 (MDI!), MDI12423, Mohd. Nor-
faizal, Amin Asyraf, Dome Nikong, Muhamad Ikhwan,
Edward Entalai & Anuar Rasyidi [holotypus, MDI].
Diagnosis
Nepenthes domei diers from N. benstonei in several lower
pitcher characteristics that include thick-leathery coria-
ceous texture (vs. coriaceous texture), broad rounded for-
mation in the lower 1/3 or 1/2 part of the lower pitcher
(vs. broad ovoid – ovoid in the lower part), two fringed
wings 5.5-7 mm wide narrowest at the base and widest
at the mouth (vs. less than 4mm runs the whole length of
the pitcher body), multicellular fringe elements up to 2-4
mm long (vs. overally shorter, 1-3 mm long). e lower
pitchers of the new species frequently burrow into humus,
a feature that has not observed in N. benstonei from Bukit
Bakar. e lower pitchers exhibit whitish red to dark ruby
red colouration with the majority of individuals sparsely
overlain with or entirely absent of specks on the inner and
outer wall (vs. brownish red-dark red, inner wall heav-
ily overlain with specks). e inorescence rachis length
28-35 cm (vs. less than 30-31 cm). Leaves and stems with-
out waxy cuticle (vs. covered with waxy cuticle).
Description
Terrestrial climber to 3.5 m tall. Stem and climbing stems
slight rounded in cross section, up to 0.7-0.9 cm in diam-
eter. Internodes 4-8 cm on climbing stems and 1.3-2.5 cm
on rosette stems. Leaves of lower stem coriaceous, lan-
ceolate, sessile, 12-13 cm long and 4-5 cm wide, sheath-
ing the stem up to ¼ of its circumference. Leaf blades
of the climbing stems narrowly linear-lanceolate, ses-
sile, 25-32 cm long, 5-8 cm wide, base contracted into
the stem up to ½-¾ of the stem, apex narrow acuminate,
tendril insertion simple. Longitudinal nerve 1-2 on each
side of the midrib, pennate nerves spreading towards
the leaf margins. Leaves of climbing stems with slight
dierentiation to those of the lower stems which is big-
ger in size. Tendrils coiled, up to 13-37 cm long. Rosette/
lower pitchers up to 15-17 cm long, 4.4-6.5 cm wide;
thick-leathery coriaceous texture; arising slight gradually
from the tendril; broad rounded formation at the lower
part ca. 1/3 with slight pronounced hip, straight cylindri-
cal towards above, widening slightly towards the mouth.
Inner surface of the portion below the lid throughout
surfaces above the hip smooth and waxiness, light green
in colour; two fringed wings up to 7 mm wide (narrow-
est at the base and widest at the mouth), run from the
top of the pitcher to the bottom of the pitcher, consist of
multicellular fringe elements up to 2-4 mm long; mouth
ovoid-rounded and oblique, concave, rising at the rear of
the pitcher towards the lid; peristome slight cylindrical,
front up to 6 mm wide, sides up to 4 mm, up to 1.2 cm
wide at the rear; outer surface with minute unicellular
trichomes, lid ovate, up to 2.7 cm long and 2.4-2.5 cm
wide, base generally rounded (not cordate), no hairs or
appendages observed on the lower surface. ~60-87 dot-
ted - rounded nectar glands scattered on the lower part
of the lid, and concentrated at the base of the lid. Upper
pitchers 10-12 cm long, 1.5-4 cm wide, thin coriaceous;
arising gradually from the tendril; slightly infundibular
to cylindrical in the lower part, upper part cylindrical
and slight widening towards the mouth; peristome simi-
lar to the lower pitchers, diers in mouth shape (wide
and length of the mouth and peristome); lid narrow
ovate, up to 1.8 cm long and 2.3 cm wide, base rounded
(not cordate), no hairs or appendages on the lower sur-
face of the lid; spur simple, unbranched less than 0.5-0.6
cm. Male inorescence a raceme, peduncle up to 13 cm,
rachis up to 35 cm, partial peduncles single-owered,
bracts absent, pedicel 11-14 mm long, tepal wide ovate,
up to 4-6 mm, staminal column 3-4 mm long, anther
head 2-2.5 mm. Colour of living plants lower pitcher
whitish red to dark ruby red throughout or green-red-
dish, sometimes with a few red-purple blotches on the
inner surfaces and lid, peristome dark red to green with
varying degrees of green and red. Leaves and stems with-
out waxy cuticle. Stems reddish-green to green. Leaves
dark green. Colour of dried specimens dark brown (lam-
ina, stem and pitcher).
Etymology
Named in honour of the rst person who discovered the
population, Mr. Dome Nikong.
Distribution
Nepenthes domei is currently known only from the
restricted area at the type locality in Setiu, Terengganu.
Ecology
Terrestrial climber in humus rich soil, immature and
mature lower/rosette pitchers tend to burrow in the soil
rich with humus up to 1/3 – 1/2 of the pitcher. Upper
hill forest habitats, at an altitude of c. 850-1000 m.
Conservation notes
Nepenthes domei and N. latiana are terrestrial climb-
ers; with N. latiana observed growing as understorey
11
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 3. Nepenthes domei, in situ: (A) Unearthed rosette pitchers with rosette leaves. (B) Immature rosette pitcher pulled out from humus
rich soil. (C) Upper pitcher. (D) Lid and inner surface of lower pitchers. (E) Leaves from lower stem. (F) Rosette/lower pitchers almost
entirely buried in the soil. (G) Habit, showing stem and leaves. Photographs by Mohd Norfaizal Ghazalli and Amin Asyraf Tamizi.
12 Mohd Norfaizal Ghazalli et al.
Figure 4. Line drawing of Nepenthes domei: (A) Habit, with inorescence. (B) Leaf blade. (C) Upper pitcher. (D) Upper pitcher, viewed
from side with spur. (E) Lower pitcher. (F) Male ower. (G) Adaxial and abaxial point view of the lid. Based on 12423 (MDI). All drawn by
Khalid Hashim.
13
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
plants on a hill slope with rocky substratum at an eleva-
tion about c. 1100 m. e vegetation of the locality con-
sisted of various montane species dominated by Dicrano-
pteris linearis (Gleicheniaceae), Dipteris conjugata (Dip-
teridaceae) and Machaerina maingayi (Cyperaceae),
thriving together with rattans, Pandanus sp. and Lepto-
spermum avescens. Some climbing stems of N. domei
were also observed scrambling over Pandanus tree.
Based on our observation, more than ve individuals
of N. domei were seen at three scattered locations, while
only one to two individuals of N. latiana were record-
ed at a single location. Both species are known to be
restricted in a single mountain since it was discovered.
us, we assign N. domei and N. latiana in this article
as Vulnerable under criterion - D2 of IUCN (2001). e
habitat had experienced logging in the past; hence, this
could explain the limited number of the individuals dis-
covered in the area.
Additional specimens examined
MALAYSIA, Kelantan, Machang, Bukit Bakar, 27.9.2006.
Yao, T.L. et al. FRI53169 (N. benstonei); Malaysia,
Terengganu, Gunung Tebu, 31.5.1974. Mohd. Shah et al.
FR I13147 (N. benstonei); Malaysia, Terengganu, Gunung
Padang, 21.3.2010. Mohd Hairul, M.A. et al. FRI 70887
(N. sanguinea); Malaysia, Pahang, Cameron Highlands,
Ulu Bertam FR, 26.4.2017. Imin, K. et al. FRI87122 (N.
macfarlanei).
DISCUSSION AND ADDITIONAL TAXONOMIC NOTES
1) Anatomical descriptions
Nepenthes domei
Holotype: MDI12423
Leaves anatomy lamina thickness 292.48-315.76
µm (Figure 9G), Hypodermis 1-3 layers on adaxial,
thickness 41.52-55.37 µm, 1 layer on abaxial, thickness
39.56-49.73 µm. Mesophyll 227.38-239.78 µm; palisade
parenchyma 1-3 layers with total 157.29-163.21 µm thick;
spongy parenchyma total 74.85-89.63 µm, with plenty
extracellular space in between of spongy parenchyma
(Figure 9G). Midrib outline Type I (Figure 9A), 267.89-
290.66 µm thick, bigger towards apex leaves; vascular
system Type I, sclerenchyma ring not observed, 482.64-
515.16 µm diameter, broader towards leaves apex; vas-
cular bundles (8 VB) (Figure 9A & B); 21.28-35.23 µm
diameter; midrib epidermis single layer, 11.35-15.45 µm
Table 1. Morphological comparison between Nepenthes domei, N. latiana, N.benstonei and N. sanguinea (Figure 1-4).
Character N. domei N. latiana N. benstonei N. sanguinea
Altitudinal range 850-1000 m 1000-1100 m 800-1350 m 300-2000 m
Habitat
Terrestrial in rich humus
soil, rosette/lower pitchers
tend to burrow in the soil
up to ½ of the pitcher body.
Terrestrial climber in upper
montane forest.
Terrestrial in secondary
vegetation and hill slopes.
Terrestrial in shrubland,
rocky areas (eg. inland
clis, mountain peaks),
forest/terrestrial.
Tendrils of the rosette and
climbing leaves
Slight coiled, up to 13-37
cm long.
With slender coiled
formation in the middle; up
to 10-14 cm long, covered
with minute hairs,
Slightly coiled, less than 60
cm long.
Tendrils slight coiled, less
than 30 cm.
Colour of lower pitchers
Whitish red to dark ruby
red with a few red-purple
blotches on the inner
surfaces and lid.
Green-yellowish-brown
hues with several red
blotches on the 1/3 of the
upper part of the pitcher
body.
Reddish-brown.Brown to reddish.
Lid of lower pitchers Narrow ovate;
lid base rounded.
Rounded; lid base retuse-
cordate. Ovate; lid base sub-ovate. Generally ovate; base
usually slight cordate.
Upper pitchers Up to 10-12 cm high × 1.5-
4 cm wide.
Up to 20-23.5 cm high ×
5-7.5 cm wide.
Less than 15 cm high × and
3 cm wide.
Less than 35 cm high × less
than 7 cm wide.
Colour of upper pitchers Green. Light green. Pale green.
Slight yellowish – green,
with several blotches on 1/3
upper part of the pitcher.
Lid of upper pitchers Narrow ovate to elliptical. Rounded. Ovate. Generally ovate, usually
slight cordate at the base.
Colour of dried specimen Dark brown. Pale brown. Brown. Brown.
14 Mohd Norfaizal Ghazalli et al.
at the abaxial and 13.53-20.53 µm at the adaxial, irregu-
lar between each other, stomata not observed; trichome
or appendages not observed (Figure 9A-F); crystal sand
and solitary sand crystals occurring in parenchyma
cells of the midrib (Figure 9C & D). Margin bluntly
pointed below, 75-80 degree pointed below, with epider-
mis thickness 29.57 µm, margin thickness 139.21 µm,
overall blade thickness 291.02-284.75 µm (Figure 9H).
Adaxial epidermis straight – straight to slightly curve,
crystal sand not observed, 2.8-4.5 µm thick; epidermis
1 layer, 2-4 sides, 14.6-29.4 µm × 7.7-27.9 µm; stomata
absent; sessile glands rounded up to 7 cells, rounded, tri-
chome absent (Figure 9M). Abaxial epidermis straight –
straight to slightly curve, 2.2-3.8 µm thick; epidermis 1
layer, irregular, more than 4-5 sides, 17.6-18.3 µm × 5.6-
19.5 µm; stomata anomocytic/ranunculaceous (Figure
9L, M & N), sessile glands rounded to ower-like in 7-8
cells, orbicular to elliptical, 42.3-59.6 µm × 40.2-51.8 µm;
trichome absent (Figure 9L). Marginal venation incom-
plete type, uni-veinlet (Figure 9I). Areolar/lamina vena-
tion incomplete (Figure 9J), type of veinlets simple, uni-
veinlet (linear-curved) (Figure 9K).
Nepenthes latiana
Holotype: MDI12424
Leaves anatomy lamina thickness 788.68 -794.50
µm (Figure 10E & H), Hypodermis 1-2 layers on adaxial,
thickness 152.89-162.33 µm, 1 layer on abaxial. Meso-
phyll 296.56-390.76 µm; palisade parenchyma 2-3 layers
342.44-387.64 µm thick; spongy parenchyma 564.59-
576.54 µm, with plenty extracellular space in between of
spongy parenchyma (Figure 10F & H). Midrib outline
Type IV, adaxial with slightly at and abaxial prominent
broad ‘U’ curve (Figure 10A), 669.63-689.45 µm thick,
smaller towards apex leaves; vascular system Type VI,
schlerenchyma ring orbicular upwards, slightly curved
at adaxial, outer ring present, central or medullary vas-
cular bundle present (3 VB), additional vascular bundle
present (several smaller size vascular bundles observed
near abaxial epidermis, 380.23-395.56 µm diameter (Fig-
ure 10A-D); up to 50.67 µm diameter; irregular between
each other; stomata not observed; trichome appendages
not observed (Figure 10A); crystal sand - miniature soli-
tary crystals occurring in parenchyma cells of the midrib
(Figure 10B & D). Margin sharp pointed pointed below
up to 85 degree, with epidermis thickness 13.35-21.58
µm, margin thickness 136.78-146.75 µm, overall blade
thickness 335.65-380.76 µm (Figure 10F). Adaxial epider-
mis straight – straight to curve, druses observed on the
adaxial surface, 3.7-6.5 µm thick; epidermis 1 layer, 4-5
sides, 15.1-38.7 µm × 5.9-22.3 µm; stomata absent; sessile
glands irregular in rounded up to 6 cells, elliptical-orbic-
ular, trichome absent (Figure 10L). Abaxial epidermis
straight – straight to curve, 2.4-4.1 µm thick; epidermis 1
layer, irregular, more than 4 sides, 14.32-28.54 µm × 43.5-
32.6 µm; stomata anomocytic/ranunculaceous (Figure
10M &N), rounded, 108.25-122.645 µm × 118.1-119.4 µm,
sessile glands not observed, trichome absent. Marginal
venation closed type, none veinlet (Figure 10K). Areolar/
lamina venation incomplete (Figure 10I), type of veinlets
simple uni-bi veinlet (curved) (Figure 10J).
Nepenthes benstonei
Leaves anatomy lamina thickness 252.96-266.43 µm
(Figure 11G), Hypodermis 1-2 layers on adaxial, thick-
ness 25.67-35.94 µm, 1 layer on abaxial, thickness 16.26-
27.38 µm. Mesophyll 159.17-172.86 µm; palisade paren-
chyma 1-3 layers with total 67.65-84.15 µm thick; spongy
parenchyma total 106.97-111.25 µm, with plenty extra-
cellular space in between of spongy parenchyma (Figure
11G & H). Midrib outline Type I, adaxial very slightly
concave, abaxial prominent ‘U-V’ Shape (Figure 11A),
380.81-391.71 µm thick, vascular system Type VI, outer
ring present (rectangular in shape), central vascular
bundle present (3 vascular bundles), additional vascular
bundle present (several smaller size vascular bundle near
abaxial epidermis) and 2 vascular bundle present on the
adaxial side of outer ring vascular bundle at the le and
right side, midrib epidermis single layered, 13.26-17.83
µm at the abaxial and 15.59-16.42 µm at the adaxial,
irregular between each other; stomata not observed; tri-
chome appendages not observed (Figure 11A & I); crys-
tal sand - solitary sand crystals occurring in masses
specically in parenchyma cells of the midrib (Figure
11B-D), druses observed (Figure 11E), starch grains
observed. (Figure 11B). Margin sharply pointed below
20-25 degree, with epidermis thickness 8.81-13.25 µm,
margin thickness 68.51-82.83 µm (Figure 11H). Adaxial
epidermis straight to wavy, crystal sand observed on the
adaxial surface, 3.5-4.9 µm thick; epidermis 1 layer, 4-6
sides, 15.6-34.5 µm × 8.4-9.3 µm; stomata absent; sessile
glands irregular in ower-like in 6-7 cells, elliptical, tri-
chome absent (Figure 11J). Abaxial epidermis sinuous,
2.5-3.8 µm thick; epidermis 1 layer, irregular, more than
4 sides, 18.42.87 µm x 6.5-36.9 µm; stomata anomocyt-
ic/ranunculaceous (Figure 11K), kidney-like, trichome
absent. Marginal venation closed type, none veinlet
(Figure 11M). Areolar/lamina venation incomplete (Fig-
ure 11L), type of veinlets simple veinlet (linear-curved)
1-2 branched (Figure 11L).
15
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Nepenthes sanguinea
Leaves anatomy lamina thickness 224.20-235.23 µm
(Figure 12E), Hypodermis 1 layer on adaxial, thickness
20.54-37.65 µm, 1 layer on abaxial, thickness 29.15-32.52
µm. Mesophyll 145.87-166.45 µm; palisade parenchyma 2
layer with total 53.06-63.32 µm thick; spongy parenchyma
total 90.71-94.99 µm, with plenty extracellular space in
between of spongy parenchyma (Figure 12E & F). Mid-
rib outline Type I, adaxial slightly attened and abaxial
arched, (Figure 7a), 383.37-445.89 µm thick, vascular
system Type VI, sclerenchyma ring orbicular upwards,
slightly curved at adaxial, outer ring present, central or
medullary vascular bundle present (4 VB), 1 additional
vascular bundle present (smaller size vascular bundles
observed near abaxial epidermis), 367.11-413.32 µm diam-
eter (Figure 7A-C); diameter up to 40.35 µm; irregu-
lar between each other; stomata not observed; trichome
appendages not observed (Figure 12A); crystal sand - soli-
tary crystals and druse occurring in parenchyma cells
of the midrib (Figure 12B-D). Margin sharp tip pointed
below up to up to 80 degree, roughly crease surface, with
epidermis thickness 6.85-9.41 µm, margin thickness
86.76-92.23 µm, overall blade thickness 89.05-106.97 µm
(Figure 12F). Adaxial epidermis straight – slight curve,
druses not observed on the adaxial surface, 3.8-7.8 µm
thick; epidermis 1 layer, 3-5 sides; stomata absent; ses-
sile glands irregular in rounded up to 5 cells, elliptical,
trichome absent (Figure 12I). Abaxial epidermis straight
– slight curve, 2.4-4.9 µm thick; epidermis 1 layer, irregu-
lar, more than 4-6 sides; stomata anomocytic (Figure 12J),
95.47-117.74 µm × 105.54-147.65 µm, sessile glands not
observed, trichome absent. Marginal venation incomplete
type, uni-veinlet (Figure 12H). Areolar/lamina vena-
tion incomplete (Figure 12I), type of veinlets – simple uni
veinlet (slight curved) (Figure 12I).
Table 2. Comparison of leaf micromorphological observation between Nepenthes domei, N. latiana, N. benstonei and N. sanguinea (Figure
5-8).
N. domei N. latiana N. benstonei N. sanguinea
Cuticular striation Adaxial: smooth-coarse;
anticlinal wall not clear;
periclinal wall sunken,
cuticular striation not
obscure and both anticlinal
and periclinal walls cannot
be dierentiated clearly.
Epidermal cell outline
obscured by appendages.
Abaxial: smooth surface,
with granular waxes; both
anticlinal and periclinal walls
cannot be dierentiated
cl early.
Adaxial: coarse; anticlinal
wall clear; periclinal wall
sunken, cuticular striation
obscure and both anticlinal
and periclinal walls cannot
be dierentiated clearly.
Epidermal cell outline
obscured by high density of
waxes.
Abaxial: coarse; with
granules waxes; both
anticlinal and periclinal walls
can be dierentiated clearly.
Adaxial: coarse; anticlinal
wall not clear; periclinal wall
sunken, cuticular striation
obscure and both anticlinal
and periclinal walls can be
dierentiated. Epidermal cell
outline obscured by high
density of waxes.
Abaxial: coarse; with aked
waxes; both anticlinal and
periclinal walls cannot be
dierentiated clearly due to
high density of akes and
appendages.
Adaxial: coarse; anticlinal
wall not clear; periclinal
wall sunken, both anticlinal
and periclinal walls can
be dierentiated clearly.
Epidermal cell outline not
obscured by high density of
waxes.
Abaxial: medium coarse;
both anticlinal and periclinal
walls can be dierentiated.
Epicuticular waxes i. Granules
present on both adaxial and
abaxial surfaces, tubular wax
on abaxial.
i. Flakes (adaxial)
ii. Granules (abaxial)
i. Flakes (adaxial)
ii) Granules (abaxial)
Flakes present on both
adaxial and abaxial
epidermises.
Stomata features Amphistomatous, sunken,
scattered mainly on the
abaxial surface, sparsely
distributed on the adaxial
surface, rounded shape.
Hypostomatic, supercial,
restricted to and sparsely
scattered on the abaxial
epidermis
surface, broad oval shape.
Hypostomatic, supercial,
restricted to and sparsely
scattered on the abaxial
epidermis
surface, broad oval shape.
Amphistomatous, sunken,
scattered mainly on the
abaxial surface, sparsely
distributed on the adaxial
surface, rounded-oval shape.
Types of trichomes i. Simple unicellular (long,
pointed tip, with slight
echinate ornamentation).
ii. Capitate glandular
trichome (multicellular
terminal).
iii. Capitate glandular
trichome (unicellular
terminal).
iv. Peltate, Scale trichome.
i. Simple unicellular (long,
pointed tip, slight echinate
ornamentation.
ii. Simple unicellular short,
blunt tip, without echinate
ornamentation
iii. Capitate glandular
trichome (multicellular
terminal).
i. Simple unicellular (long,
pointed tip, smooth).
ii. Capitate glandular
trichome (unicellular
terminal).
iii. Capitate glandular
trichome (multicellular
terminal).
i. Capitate glandular trichome
(multicellular terminal).
16 Mohd Norfaizal Ghazalli et al.
Results of the study revealed a number of interest-
ing features with some characters which could serve
as taxonomic and diagnostic value, while assist in giv-
ing additional evidences in describing N. domei and N.
latiana against related species. Anatomical diagnos-
tic characters that can be used in directly dierentiat-
ing and possessed taxonomic value are sclerenchyma
ring type and shape, and marginal venation and lamina
venation that enumerate signicant taxon separation
between N. domei and N. benstonei, and N. latiana
and N. sanguinea. Supportive micromorphology and
anatomy characters which are also useful in assisting
these taxa identication (diagnostic characters) in new
taxon descriptions are noted as below, with identica-
tion key of the species:
2) Diagnostic characters of Nepenthes domei and N. latif-
ana based on leaf anatomy and micromorphology
Anticlinal walls of lamina epidermal cells under light
microsope (LM)
According to Stace (1969), the lamina surface has
been the focal subject of investigations (as compared
with other plant surfaces) that some of studied taxa
showed consistent characteristics which oen serve as a
taxonomic application. From observations of the lamina
surface in studied Nepenthes, the anticlinal wall of N.
domei is straight to slight curve for both abaxial and
adaxial epidermis surfaces while N. benstonei showed
straight to wavy adaxial epidermis surface and sinuous
abaxial epidermis surface. On the other hand, N. lati-
ana showed straight to straight curve epidermis wall for
both of its surfaces as compared to N. sanguinea which
possessed straight – slightly curve and straight for its
abaxial epidermis. erefore, these characters appear to
be diagnostic for those two new species, respectively.
Margin and Midrib Transverse Section (TS)
Studied Nepenthes species showed interspecies vari-
ations in lamina margin transverse section (TS) which
can serve as a useful characteristic for species identi-
cation. e outline of the TS of the margin is bluntly
rounded (pointed 75-80°) for N. domei, sharply pointed
downwards that reached 85° for N. latiana, sharply
pointed with 20-25° curve for N. benstonei, while N. san-
guinea having up to 80° and rough creased abaxial and
adaxial surfaces. Midrib outline for N. domei, N. san-
guinea and N. benstonei is characterised as Ty p e I (abax-
ial with prominent U-V shape and adaxial with very
Table 3. Comparison of leaf anatomical features between Nepenthes domei, N. latiana, N. benstonei and N. sanguinea (Figure 9-12).
N. domei N. latiana N. benstonei N. sanguinea
Leaf margin Bluntly pointed below 75-80°.Sharply pointed below up
to 85°.
Sharply pointed below
20-25°.
Sharply pointed below up to
up to 80°, with roughly crease
surface.
Midrib Outline Type I. Outline Type IV. Outline Type I. Outline Type I.
Vascular bundle
arrangement
Arrangement Type I. Arrangement Type VI. Arrangement Type VI. Arrangement Type VI.
Marginal venation Incomplete type, uni-veinlet. Closed type, none veinlet. Closed type, none veinlet. Incomplete type, uni-veinlet.
Areolar venation Incomplete, type of veinlets
simple, uni-veinlet (linear-
curved).
Incomplete, type of veinlets
simple uni-bi veinlet
(curved).
Incomplete; type of veinlets
simple veinlet (linear-curved)
1-3 branched.
Incomplete, type of veinlets-
simple uni-veinlet (slightly
curved).
Pattern of anticlinal
cells
Adaxial: straight – straight to
slightly curved.
Abaxial: straight – straight to
slightly curved.
Adaxial: straight – straight to
curved.
Abaxial: straight – straight to
curved.
Adaxial: straight to wavy,
crystal sand observed on the
adaxial surface.
Abaxial: sinuous.
Adaxial: straight –slightly
curved.
Abaxial: straight – slightly
curved.
Stomata feature Anomocytic (type of
stomata-without subsidiary
cells);
Homostomatic (only one
type of stomata present);
Amphistomatous (stomata
present on both epidermis
surfaces).
Anomocytic; Homostomatic;
Hypostomatic (stomata
only present on abaxial
epidermis).
Anomocytic; Homostomatic;
Hypostomatic.
Anomocytic; Homostomatic;
Amphistomatous.
17
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
slight concave shape) while N. latiana having Ty pe
IV (abaxial with prominent V shape and adaxial with
prominent wide V shape) midrib outline. For midrib TS,
the variations are as follows: N. domei is having Ty p e I
(adaxial vascular bundle in stacks, medullary vascular
bundle absent, additional vascular bundle present in sev-
eral smaller size nearby the abaxial epidermis) vascular
bundle arrangement, while N. latiana, N. benstonei
and N. sanguinea are having Ty pe V I vascular bundle
arrangement (outer ring present, elliptic; central med-
ullary vascular bundle present in scattered formation;
additional vascular bundle present in several smaller size
nearby the adaxial side of outer ring at the le and right
side). ese observations (disparity in vascular bundle
arrangement combined with midrib outline) agree with
some studies conducted by Metcalfe and Chalk (1950)
who demonstrated the two characteristics could greatly
serve for diagnosis in certain species, as recorded and
showed in some Parashorea species.
Marginal and areolar venation
e variation of leaf venation patterns of angio-
sperms was extensively studied and classied by Hickey
(1979) and Sun et al. (1997) and this further addresses
the importance of comprehensive treatment in taxon
dierentiation. Leaf architecture is primarily used for
classication which includes the leaf shape, leaf mar-
gin structure and other possible characters. Indeed, the
characterisation of an angiosperm leaf venation pattern
starts by observing the primary veins and then pro-
ceeded to the branching secondary veins. Combination
of marginal venation and areolar venation patterns for
every taxon are unique, even for those taxa classied
under the same genus (Inamdar et al. 1983). In studied
Nepenthes species, marginal and lamina/areolar vena-
tion features showed diagnostic characteristics that are
useful in species dierentiation and evaluation. Nepen-
thes domei and N. sanguinea showed incomplete uni-
veinlet, while N. benstonei and N. latiana are charac-
terised with closed type, with none veinlet. Another fea-
ture analysed is the areolar / lamina venation, in which
all taxa showed incomplete areolar venation but varied
in type of venation ending as follows: N. domei showed
simple veinlet with uni-veinlet (linear to curved), while
N. latiana and N. benstonei shared the same feature of
uni-bi veinlet (curved) that slightly diers in N. bensto-
nei venation ending with linear-curved (1-3 branched)
and N. sanguinea characterised with uni-veinlet (slight
curved). ese variable patterns of leaf venation are tax-
onomically signicant in these pitcher plants species as
they can be used as additional evidences for species dif-
ferentiation.
Cuticular wax and epidermal sculpturing under scan-
ning electron microscopy (SEM)
Cuticular sculpturing also holds considerable diag-
nostic values as reported by Wilkinson (1979) and Wu et
al. (2005). In this study, the epidermal surfaces revealed
a number of important micromorphological characters,
exhibited interesting interspecies variations that are sig-
nicant for species identication. Leaf epidermis sur-
faces (abaxial and adaxial) of the investigated Nepenthes
species that were viewed under the SEM (low to high
magnications) gave certain features – distribution of
idioblastic elements such as trichomes, glands and sto-
mata, combined with the appearance of epicuticular
wax. In this study, the appearance of epicuticular wax
on both of the leaf surfaces could be tubular granular
or aked, or a combination of dierent types of wax.
Nepenthes domei diered from others for having gran-
ular wax on both abaxial and adaxial epidermis sur-
faces, and tubular wax only on abaxial surface (Figure
5). Meanwhile its closely related species, N. benstonei,
showed numerous wax akes (Figure 7) for its adaxial
epidermis surface and granules for its abaxial surface.
It seems the presence of these numerous akes sup-
ports Clarke’s (2001) observation saying that the leaves
and stems of N. benstonei were covered with thick waxy
cuticle. Nepenthes latiana showed similar features as
that of N. benstonei, while N. sanguinea had wax akes
on both abaxial and adaxial surfaces. Stomata features
also implicate a signicant diagnostic value for our spe-
cies of interest. Nepenthes domei and N. sanguinea leaves
are amphistomatous (stomata on both surfaces); how-
ever N. benstonei and N. latiana are hypostomatous
(stomata only on the lower surface). While hypostoma-
tous stomata feature is common for the majority of spe-
cies for monocotyledons and dicotyledons (Meidner and
Manseld 1968), amphostomatous feature is much rare
(Drake et al., 2018). Drake et al. (2018) further points
out that amphistomaty may help in better CO2 absorp-
tion but with higher rate of transpiration and this dis-
covery of two dierent groups of leaves (hypostomatous
vs. amphostomatous) within the same genus is truly
intriguing. The indumentum feature in dicotyledons
was reported to consist of simple unicellular and vari-
ous glandular trichomes (Metcalfe and Chalk, 1950). In
this study, glandular trichomes was present in all spe-
cies but N. domei showed diagnostic character with the
observation of scale-type trichomes, known as peltate,
18 Mohd Norfaizal Ghazalli et al.
on its abaxial epidermis surface that did not exist in
other investigated species. Simple unicellular trichomes
are common and present in all species of studied Nepen-
thes except in N sanguinea that also can be postulated
as a diagnostic characteristic that dierentiates N. latif-
ana from N. sanguinea. is trichome type can be long
or short, and has either thick wall ornamentation or
smooth.
Identication key based on anatomy and micromorphol-
ogy characters
1 a. Midrib outline Type I; vascular bundle arrangement
Type I, with peltate trichome ...................................N. domei
b. Midrib outline Type IV; vascular bundle arrangement
type VI ...................................................................................... 2
2 a. Margin transverse section sharply pointed over 20-25° 3
b. Margin transverse section sharply pointed below up to
85° ..........................................................................N. latiana
3 a. Adaxial epidermal cell outline obscured by high density
of waxes ................................................................ N. benstonei
b. Adaxial epidermal cell outline not obscured by high
density of waxes.......................................................................4
4 Simple unicellular trichome absent, only peltate glandular
trichome (multicellular terminal) observed ...N. sanguinea
3) Molecular analysis of ITS sequence to infer the relation-
ship between N. benstonei and other species
e ITS is a nuclear DNA region which is highly
conserved and it has been used successfully to discrimi-
nate dierent plant species including Nepenthes (Giudi-
celli et al. 2015; Gogoi and Bhau 2018). Based on nucleo-
tide search on the public NCBI database (BLASTn), we
noted partial ITS region (~610bp) is more than 95%
conserved among the majority species in the Nepenthes
genus and the ITS may not be sensitive to dierentiate
sub-populations under the same species. However, we
tested the degree of ITS sequence conservation from a
few dierent populations of N. sanguinea in Peninsu-
lar Malaysia and it is proven the partial ITS sequence
is 100% conserved within this particular species (data
not shown), saying that ITS intraspecic variation of a
closed population should be close to 0.
In order to further establish N. domei as a separate
taxon from its most closely related species N. bensto-
nei, we conducted a molecular analysis by comparing
the partial ITS sequence from both plants (Figure 13)
and then inferred phylogenetic relationship (NJ meth-
od) of the new species with nine peninsular Nepen-
thes species (Figure 14). Based on sequence alignment,
N. domei sp. nov shares up to 97.2% ITS nucleotide
identity to N. benstonei, and the 2.8% nucleotide dis-
similarity – a gure signicant enough to dierentiate
Nepenthes species in this study – is due to nucleotide
polymorphisms that occur at 15 dierent positions,
denoted as non-coloured letters (Figure 13). In addi-
tion to this, the phylogenetic inferring has positioned
N. domei on a separate branch from N. benstonei in
the highland/intermediate highland clade (Figure 14).
is molecular scrutiny – on top of morphology, leaf
anatomy and micromorphology – serves as an addition
evidence that has positioned N. domei as a distinct
species from N. benstonei.
CONCLUSION
e combination approach of morphology-anato-
my-micromorphology analyses used in this comparison
study is useful in providing as much aspects as possible
in distinguishing plant characteristics of N. domei and
N. latiana against their congener species, N. bensto-
nei and N. sanguinea. As a whole, pitcher morphology
of both new species combined with detailed descriptions
of the midrib, lamina and epidermis have addressed
comprehensive taxonomic resolution in dening these
new Nepenthes species. Also, the phylogenetic inference
has provided an additional supportive evidence for the
placement of N. domei as a new species. Hence, N. domei
and N. latiana described in this paper are condently
positioned as two new species discovered from eastern
part of Peninsular Malaysia.
ACKNOWLEDGEMENTS
is study was carried out under Forestry Depart-
ment of Peninsular Malaysia (Ministry of Energy and
Natural Resources) permit ref. JH/100 Jld. 22 (41). We
are grateful to Dr. Richard Chung Cheng Kong (For-
est Research Institute Malaysia, FRIM) for his guidance
and help in obtaining permission involving herbarium
records and observation of the pitcher plants herbarium
specimens (KEP). Special thanks go to Madam Salman-
iza Salleh and Miss Nurshahidah Mohd Rusli for their
technical assistance in the anatomical and palynological
works as well as in herbarium curations. We also extend
our gratitude to Muhammad Ikhwan Afandi Md Daud
for his assistance during the eld work.
19
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 5. Nepenthes domei: (A) Abaxial epidermis; stoma (red arrow) and peltate trichome (yellow arrow). (B) Stomata and epicuticular
wax. (C) Stoma (arrow). (D) Peltate (scale) trichome. (E) Adaxial epidermis with tubular wax. (F) Stoma deep embedded into tubular wax
(arrow) observed on the adaxial epidermis. (G) Granular wax mixed with tubular. (H) Cuticular striation. SEM images by by Mohd Nor-
faizal Ghazalli and Ahmad Zaki Zaini.
20 Mohd Norfaizal Ghazalli et al.
Figure 6. Nepenthes latiana: (A) Abaxial epidermis adorned with numerous stomata. (B) Abaxial stomata (arrow). (C) Stomata (arrow) on
abaxial surface with wax granules. (D) Peltate glandular trichome. (E) Peltate glandular trichomes (yellow arrows) and stoma (red arrow). (F)
Adaxial epidermis with wax akes but without stomata. (G) Capitate glandular trichome (arrow). (H) Simple, unicellular trichome, long pointed
(arrow). (I & J) Variation of multicellular peltate glandular trichome (arrow). SEM images by Mohd Norfaizal Ghazalli and Ahmad Zaki Zaini.
21
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 7. Nepenthes benstonei: (A & B) Abaxial epidermis with stomata (arrows). (C) Abaxial stoma surrounded by wax granules (arrow).
(D) Simple, unicellular trichome, long pointed (arrow). (E) Adaxial surface absent of stomata. (F) Capitate glandular trichome (arrow) sur-
rounded by wax akes. (G) Multicellular peltate glandular trichome (arrow). (H) Flakes observed on the adaxial epidermis. SEM images by
Mohd Norfaizal Ghazalli and Ahmad Zaki Zaini.
22 Mohd Norfaizal Ghazalli et al.
Figure 8. Nepenthes sanguinea: (A) Abaxial epidermis. (B) Stomata (arrow). (C) Stoma. (D) Adaxial epidermis with stoma (red arrow) and
peltate glandular trichome (yellow arrow). (E) Adaxial epidermis surface. (F) Stoma. (G & H) Variation of multicellular peltate glandular
trichomes observed on the adaxial epidermis surface. SEM images by by Mohd Norfaizal Ghazalli and Ahmad Zaki Zaini.
23
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 9. Nepenthes domei: (A) Midrib TS. (B, C & D) Solitary sand crystal (arrows). (E) Parenchyma cells. (F) Simple, unicellular trichome
(arrow). (G) Lamina TS. (H) Margin TS. (I) Margin venation. (J) Lamina venation. (K) Type of veinlet. (L) Adaxial epidermis showing a
stoma. (M & N) Abaxial epidermis with stomata. Photographs by Mohd Norfaizal Ghazalli and Amin Asyraf Tamizi.
24 Mohd Norfaizal Ghazalli et al.
Figure 10. Nepenthes latiana: (A) Midrib TS. (B) Solitary crystal (arrow). (C) Vascular bundle (arrow). (D) Solitary sand crystal. (E)
Lamina TS. (F) Margin TS. (G) Simple, unicellular trichomes (arrow). (H) Lamina TS. (I) Lamina venation. (J) Type of veinlet. (K) Margin
venation. (L) Adaxial epidermis. (M) Abaxial epidermis with stomata. (N) Stomata. Photographs by Mohd Norfaizal Ghazalli and Amin
Asyraf Tamizi.
25
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 11. Nepenthes benstonei: (A) Midrib TS. (B) Observation of solitary crystal (yellow arrow) and starch grain (orange arrow). (C) Soli-
tary crystal (arrow). (D) Solitary crystal. (E) Druses. (F) Vascular bundle. (G) Lamina TS. (H) Margin TS. (I) Simple, unicellular trichome.
(J) Adaxial epidermis. (K) Abaxial epidermis with stomata observation. (L) Lamina venation. (M) Margin venation. Photographs by Mohd
Norfaizal Ghazalli and Amin Asyraf Tamizi.
26 Mohd Norfaizal Ghazalli et al.
Figure 12. Nepenthes sanguinea: (A) Midrib TS. (B) Solitary crystal (arrow). (C) Solitary crystal (arrow). (D) Solitary sand crystal. (E) Lam-
ina TS. (F) Margin TS. (G) Lamina venation. (H) Marginal venation. (I) Abaxial epidermis. (J) Adaxial epidermis. Photographs by Mohd
Norfaizal Ghazalli and Amin Asyraf Tamizi.
27
Two new species of pitcher plants from Terengganu, Peninsular Malaysia
Figure 13. Pairwise alignment (optimal Global alignment) of partial ITS sequences (610 bp) from N. domei sp. nov. and N. benstonei
(AB675710.1) using BioEdit Sequence Alignment Editor. Unmatched nucleotides are not coloured and matching (conserved) nucleotides
are represented by coloured blocks.
Figure 14. Phylogenetic tree (NJ method) reconstructed using ITS sequences obtained from the public database (NCBI) and ITS isolated in
this study. Bootstrap values are as indicated above relevant branches and an American carnivorous pitcher plant (Sarracenia ava) serves as
an outgroup.
28 Mohd Norfaizal Ghazalli et al.
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Table of contents
Comments
Riccardo Maria Baldini
The impact of Covid-19 crisis on Plant Taxonomy:
will we be able to approach to plant taxonomy as
in the past? 3
Articles
Mohd Norfaizal Ghazalli, Amin Asyraf Tamizi,
Dome Nikong, Edward Entalai Besi, Muhamad
Ikhwanuddin Mat Esa, Anuar Rasyidi Mohd
Nordin, A. Latiff, Ahmad Zaki Zaini, Mohamad
Alias Shakri
Nepenthes latifana and N. domei
(Nepenthaceae), two new species of pitcher
plants from Terengganu, Peninsular Malaysia 5
Wong Sin Yeng, Peter C. Boyce
Studies on the Dipterocarpaceae of Borneo, II.
Ant stipule-brood sites and extra oral nectary
association in saplings of Shorea macrophylla
[sect. Pachycarpae] in Sarawak, Malaysian Borneo 29
Daniel Francis Brunton, Paul Clayton Sokoloff
A Review of North American Isoetes engelmannii
(Isoetaceae) complex hybrids, including the
description of I. × fernaldii, hyb. nov. and I. ×
karenae, hyb. nov. 35
Abdulwakeel Ayokun-nun Ajao, Annah
Ntsamaeeng Moteetee
Taxonomic signicance of vegetative and
reproductive morphology in southern Africa
Rhynchosia sect. Rhynchosia (Fabaceae:
Papilionoideae, Phaseoleae) 51
Lorenzo Gallo
Sempervivum guillemotii Lamotte
(Crassulaceae), a rediscovered houseleek of the
French-Italian Alps 69
Angelo Troia, Vincenzo Ilardi, Elisabetta Oddo
Monitoring of alien aquatic plants in the inland
waters of Sicily (Italy) 77
Special section on Araceae
Peter C. Boyce
Introduction 85
Wong Sin Yeng, Jyloerica Joling, Joan T.
Pereira, Alviana Damit, Peter C. Boyce, Alistair Hay
Studies on Potheae of Borneo II: Pedicellarum
M.Hotta subsumed into Pothos L., and
recognition of three new species 89
Kartini Saibeh, Peter C. Boyce, Wong Sin Yeng
Studies on Schismatoglottideae (Araceae) of
Borneo LXIX: Schismatoglottis imbakensis, a
new species of the Patentinervia Complex from
ultramac rocks, Sabah 105
Wong Sin Yeng, Peter C. Boyce
Studies on the Alocasia Schott (Araceae) of
Borneo III: Alocasia puncakborneensis, a new
species belonging to the Princeps Complex 111
Wong Sin Yeng, Peter C. Boyce, Alistair Hay
Studies on Homalomeneae (Araceae) of
Sumatera VI: Two remarkable new species of
Homalomena [Chamaecladon Clade] 117
Marco Cedeño-Fonseca, Alistair Hay, Michael
H. Grayum, Mario A. Blanco
Two new endemic species of Monstera (Araceae:
Monsteroideae: Monstereae) from Golto in
southern Costa Rica 123
Reviews
Riccardo Maria Baldini
Mats Thulin - The Genus Boswellia
(Burseraceae). The Frankincense Trees 133
WEBBIA
Journal of Plant
Taxonomy and Geography
June 2020
Vol. 75 - n. 1
