A PROTOCOL FOR FLORAL ODOUR ANALYSES OF SCHISMATOGLOTTIS

CALYPTRATA (ARACEAE: SCHISMATOGLOTTIDEAE)

NUR ADILAH BINTI MUHAMMAD ARIF (24521)

Bachelor of Science with Honours

(Plant Resources Science and Management)

2012

Faculty of Resource Science and Technology

A Protocol for Floral Odour Analyses of Schismatoglottis calyptrata

(Araceae: Schismatoglottideae)

Nur Adilah Binti Muhammad Arif (24521)

This Report is submitted in partial fulfillment of the requirements of the Degree of Bachelor of

Science with Honours (Plant Resource Science and Management)

Plant Resource Science and Management Programme

Department of Plant Science and Environmental Ecology

Faculty of Resource Science and Technology

Universiti Malaysia Sarawak

2012

i

APPROVAL SHEET

Name of Candidate : Nur Adilah Binti Muhammad Arif

Title of Dissertation : A Protocol for Floral Odour Analyses of Schismatoglottis calyptrata

(Araceae: Schismatoglottideae)

…………………………………………..

(Dr. Wong Sin Yeng)

Supervisor

……………………………………..

(Dr. Siti Rubiah Binti Zainudin)

Coordinator

Plant Science and Management Program

Department of Plant Science and Environmental Ecology

Faculty Resource Science and Technology

Universiti Malaysia Sarawak

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DECLARATION

I hereby declare that the report is based on my original work except for quotation and citations

which have been duly acknowledgment. I also declare that it has not been previously or

concurrently submitted for any degrees at UNIMAS or any other institutions of higher

learning.

……………………………

(Nur Adilah Binti Muhammad Arif)

Plant Resource Science and Management Program

Department of Plant Science and Environmental Ecology

Faculty of Resource Science and Technology

Universiti Malaysia Sarawak.

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ACKNOWLEDGEMENTS

In the name of Allah the Almighty, with grateful pleasure and profound sense of

gratitude, I, Nur Adilah Binti Muhammad Arif would like to express my most cordial and

humble thanks to my respected supervisor Dr. Wong Sin Yeng for her valuable guidance, keen

interest, inspiration, encouragement, time spent and moral support throughout completion of

my study.

Special appreciation and gratitude is also addressed to MSc candidate, Mr Hoe Yin

Chen for his useful information, guidance, materials provided and valuable suggestion in this

study. My sincere thank is also extended to my mentor, Prof. Dr. Cheksum @ Supiah Binti

Tawan for her continuous moral support and kindness. Special thanks to the lab assistants for

their kind co-operations.

Lastly, I wish to express my sincere appreciation especially to Nurul Fatini Shahirah

Suharizan, Zulaikha Zainal, Nurul Atiqah Abdul Rahman, Khairul Afiq Mohamed Noordin

and Muhammad Firdaus Latip who gave a lot of support and motivation. Also not forgotten, to

all my colleagues, who had assisted me in conducting this study either directly or indirectly.

From the deepest depth of my heart, I express my thanks to my beloved mother and

father, and also the rest of my family members who kept supporting.

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TABLE OF CONTENTS

Page

Approval Sheet i

Declaration ii

Acknowledgements iii

Table of Contents iv

List of Tables and Figures x

Abstract 1

1.0 Introduction

1.1 Problem Statements 5

1.2 Objectives 5

2.0 Literature Review

2.1 Bornean Araceae 6

2.2 Tribe Schismatoglottideae 9

2.3 Schismatoglottis 12

2.4 Schismatoglottis calyptrata complex 15

2.5 Floral Odours and Pollination Biology 19

3.0 Materials and Methods

3.1 Sampling Location 21

3.2 Floral Odour Trapping Method 22

3.3 Sampling of Floral Odours 24

3.4 Gas Chromatography 25

3.5 Mass Spectrometry Analysis 30

v

4.0 Results and Discussions

4.1 Compounds 32

4.2 Chromatograms 34

4.3 Discussions 44

5.0 Conclusions and Recommendations 54

References 56

1

A Protocol for Floral Odour Analyses of Schismatoglottis calyptrata

(Araceae: Schismatoglottideae)

Nur Adilah Muhammad Arif

Plant Resource Science and Management Programme Department of Plant Science and Environmental Ecology

Faculty of Resource Science and Technology Universiti Malaysia Sarawak

ABSTRACT

The Schismatoglottis calyptrata complex comprises about 30 species of herbs originating from the Asian humid

tropics belonging to the plant family Araceae. Inflorescences of Schismatoglottis flower by a series of functional ‘phases’, beginning with female flower activity (the ‘pistillate phase’) and ending with pollen release (‘staminate

phase’), with these phases separated by a transitional period. The production of primary floral attractants was in

the form of powerful esteric-like odours. Several species of the Calyptrata complex frequently occur together in a

habitat, with the different species often flowering at the same time yet nowhere are there recorded interspecific

hybrids. It thus seems very probable that aspects of the species’ pollination maintain distinct lineages of the

various species, even in situations of multi-species flowering. A protocol for floral odour analyses of S.

calyptrata was established in this study. The main compound of the floral scent trapped was examined by using

gas-chromatography mass-spectrometry (GCMS) techniques with the aim of identifying the compounds emitted.

Four compounds were identified and characterised, representing 73.74-93.16% of the total GC area. The major

volatile representing about 50% was the Methyl 3-methyl-3-butenoate. This compound was found in higher

concentrations from female anthesis of S. Calyptrata (F3). Generally, a significant difference in the composition

of volatile compounds between the compounds from the same antheses and from different geographic regions was recorded. The results suggest that climatic factors, latitude and longitude affect the formation of volatiles.

Key terms: Araceae – Schismatoglotideae – Schismatoglottis calyptrata –floral odour – GCMS

ABSTRAK

Kompleks Schismatoglottis calyptrata terdiri daripada kira-kira 30 spesies herba yang berasal dari kawasan

lembap tropika Asia, keluarga tumbuhan Araceae. Pendebungaan bunga Schismatoglottis oleh satu siri 'fasa'

berfungsi, bermula dengan aktiviti bunga betina ('fasa pistilat') dan berakhir dengan pembebasan debunga ('fasa

staminat'). Fasa-fasa ini dipisahkan oleh tempoh peralihan. Pengeluaran penarik utama bunga ini dalam bentuk

bau esterik yang kuat. Beberapa spesies daripada kompleks Calyptrata sering berlaku bersama-sama di dalam

habitat, dengan spesies berbeza yang sering berbunga pada masa yang sama tetapi mana yang ada direkodkan

sebagai kacukan antaraspesies. Berkemungkinan besar bahawa aspek pendebungaan spesis mengekalkan

keturunan yang berbeza daripada pelbagai spesies, walaupun dalam keadaan berbunga multi-spesies. Satu

protokol untuk analisis bau bunga S. calyptrata telah diwujudkan dalam kajian ini. Sebatian utama aroma bunga

yang terperangkap telah diperiksa dengan menggunakan teknik kromatografi gas-spektrometri jisim (GCMS)

bertujuan untuk mengenal pasti sebatian dikeluarkan. Empat sebatian telah dikenal pasti dan ciri-ciri, mewakili 73.74-93.16% daripada jumlah kawasan GC. Sebatian ruapan utama yang mewakili kira-kira 50% adalah

Methyl 3-metil-3-butenoate. Sebatian ini didapati dalam kepekatan yang lebih tinggi dari bunga putik betina S.

Calyptrata (F3). Secara umumnya, perbezaan yang signifikan dalam komposisi sebatian meruap antara sebatian

dari putik-putik bunga yang sama dan dari kawasan geografi yang berbeza telah direkodkan. Hasil kajian

mencadangkan bahawa faktor cuaca, latitud dan longitud memberi kesan kepada pembentukan meruap.

Key terms: Araceae – Schismatoglotideae – Schismatoglottis calyptrata –floral odour – GCMS

2

1.0 INTRODUCTION

The Araceae is a family of herbaceous monocots. The family members are highly

diverse in life form from submerged to free-floating aquatics. The leaves can vary

considerably from species to species. Species in Araceae are often rhizomatous or tuberous

and are often found to contain calcium oxalate crystals or raphides. Inflorescences of some

plants in this family are thermogenic (heat-producing) as the temperature of the inflorescences

can reach up to 45 degrees Celsius even when the surrounding air temperature is much lower

(Gibernau, et al. 2005). Boyce et al. (2010) reported that the total aroid flora for Borneo quite

likely exceeds 1,000 species. In short, Borneo is an aroid habitat of global significance; and

arguably one of the richest and most diverse on the planet. Barabe & Lacroix (2001) explained

the lack of floral developmental studies in the Araceae family results in part from the

difficulty in obtaining enough material to adequately document the early stages of

development. An alpha-taxonomic revision of the Malesian species, one of the results of

which was the recognition of six informal species grouping based accordingly on short

architechture and the senescence mechanics of the upper spathe (Hay & Yuzammi, 2000).

Tribe Schismatoglottideae is a predominantly Indomalayan taxon comprising 13

genera of terrestrial, lithophytic or rheophytic herbs with a primary distribution centered on

Borneo but with the total distribution ranging to South West Myanmar, South West China, and

Vanuatu (Hay & Yuzammi, 2000). Schismatoglottis Zoll. & Moritzi is the largest genus in

Schismatoglottideae with over 200 species. Spadix structure of Schismatoglottis is complex

and it is divided into a lower female flower zone and a middle male flower zone with interstice

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(sterile female and male flowers) and often with a sterile appendix. Schismatoglottis

inflorescences are generally rather small and inconspicuous, characterized by a complex

spadix with up to three different kinds of staminode present, the fertile elements segregated

into male and female zones on the spadix often separated by a sterile zone, and a sterile

appendix (Hay & Yuzammi, 2000).

These flowers presumably rely on particular pollinator species, and attract them by

combinations of floral traits that include floral odours, size and colour of floral organs,

flowering timing, and quality of the rewards (Sakai & Inoue 1999). Stebbins (1971) exclaimed

that such species-specific plant-insect interactions are interpreted as “the most effective

pollinator principle”. Mayo et al. (1997) reported the Araceae is one of the most abundant

families of tropical rainforest understory. Some species of Araceae are visited by some beetles

of the families Scarabaeidae, Staphylinidae and Nitidulidae (Gibernau, 2003). However, some

of the beetles are ineffective pollinators (Young, 1986; Gracia-Robledo et al., 2004).

Scents are more complex, as floral fragrances different very widely among species in

terms of the number, identity and relative amounts of constituent volatile compounds. Dobson

(1991) and Flamini et al. (2002) reported insects are able to distinguish between complex

floral scents mixtures and discriminate flower visits depending on floral scent contribute to

volatile production. Solid Phase Micro Extraction or “SPME” presents many advantages over

other methods allowing sampling and preconcentration in a single step (Pawliszyn, 1997;

Prosen and Zupancic-Kral, 1999; Sides et al., 2000; Kim & Lee, 2002; Augusto et al., 2003).

4

The analysis of flower volatiles requires special methods for their isolation with

enrichment. Living flowers show a continuous change in their volatile profile that depends on

genetic and external factors (Stashenko & Martinez, 2008). According to Kumano &

Yamaoka (2006), most Araceae plants have erect inflorescences (spadices) with surrounding

spathes. Scents emitted by vegetative plant parts effectively function as allelopathic agents,

herbivore deterrents or as attractants for the herbivores natural enemies, and flower volatiles

are traditionally regarded as pollinator attracting signals (Pichersky & Gershenzon, 2002;

Raguso, 2008). Insects are able to distinguish between complex floral scents mixtures and

discriminate flower visits depending on floral scent content. In addition, different parts of the

flower such as pollen and petals contribute to the production of floral odour (Dobson, 1991;

Flamini et al., 2002).

In this study, the main compound in complex mixtures of Schismatoglottis calyptrata

was examined by using gas-chromatography mass-spectrometry (GCMS) techniques with the

aim of identifying the different compounds of complex floral scents mixtures. The sample was

taken from one of the species area, Kampung Sikog, Kota Padawan, Sarawak. Initially, the

fragrances emitted by whole inflorescences were identified without any disruption from the

aroma profiles. Once the compounds had been identified, they were quantified in each of the

individual parts of excised flower which are female zone and male zones.

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1.1 Problem Statements

According to Hay & Yuzammi (2000), there are about 40 species in the

Schismatoglottis calyptrata group in Malesia. Several species of the Calyptrata complex

frequently occur together in habitat with the different species often flower at the same time yet

nowhere are there recorded interspecific hybrids. It thus seems very probable that aspects of

the species pollination maintain distinct lineages of the various species, even in situations of

multi – species flowering. The study focused on the species complex from Kampung Sikog

(Kota Padawan, Sarawak).

1.2 Objectives

The objectives of this study were as follows:

i. To establish a protocol of floral odour analyses for Schismatoglottis calyptrata

complex.

ii. To identify the main compounds of the floral odour for Schismatoglottis calyptrata.

iii. To link the odour composition of the inflorescences to the pollinators (which also

included various insect visitors).

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2.0 LITERATURE REVIEW

2.1 Bornean Araceae

The word “arum” is derived directly from ancient Greek “aron” and individual species

of the Araceae have been recorded by many botanists and historians since those ancient times

(Mayo et al., 1997). Besides, Araceae group especially with regard of vegetative structure is

the most diverse of any plant family distributed worldwide. Mayo et al. (1997) stated that

within the Araceae, the subfamily Aroideae consists of 74 genera and is characterized by the

presence of unisexual flowers. The floral ontogenetic pattern in Schismatoglottis represents a

different pathway in the evolution of unisexual flowers in the subfamily Aroideae.

Araceae group is the richest and the most diverse area is Central and South America

(Neotropic ecozone), notably along the mountainous Pacific coast of north eastern South

America. The next richest assemblage is in Southeast Asia, particularly the Malay

Archipelago. The Malay Archipelago includes the major landmasses (West to East): Sumatera,

the Malay Peninsula including Singapore, Jawa and Borneo. The Malay Archipelago forms

part of the larger informally ecological region which is known as Malesia. Malesia consists of

the Malay Archipelago, the remainder of Indonesia, almost of the Philippines, Timor Leste,

and the island of New Guinea. Malesia comprises much of the Indomalayan ecozone, although

from a species distribution standpoint it is necessary to include the western most islands of the

tropical Pacific (Oceania ecozone) as far east as Vanuatu, together with the islands to the north

and northwest of Australia. The next richest region for aroids are tropical Africa, followed by

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warm temperate Eurasia, India, Madagascar and the Seychelles, Southern Africa, and North

America including northern most Mexico (Mayo et al., 1997).

The aroids or common named as keladi belongs to Araceae family. This herbaceous

monocotyledon plants are characterized by their inflorescences, which consists of a spadix and

a spathe. The minute flower surrounded the spadix and the spathe cover the spadix before it’s

mature. There are two types of inflorescences occur in Aroids family named bisexual flowered

spadix with simple undifferentiated spathe and unisexual flower spathe divided into a limb

(blade) and convolute lower tube. The first taxonomic study of family Araceae in Peninsular

Malaysia was done by Ridley (1925). The utilization of Malaysian plants as a medicinal plants

including aroids are reported by Burkill & Haniff (1930) whereas Ghani (1983) have studied

the Aroids as sources of food and ornamental plants. Neoh (1992) has discovered Aroids that

have potential as medicinal plants. In addition, Kress (1995) has reported several medicinal

plants including Aroids. Besides, others studied are done by Jacobsen (1995), Hay (1996), and

Mansor & Sulaiman (1997).

Our knowledge of the Aroids of Borneo sometimes based in part on data and samples

gathered by many researchers with different levels of interest in the family. Borneo is an Aroid

habitat of global significance, and arguably one of the richest and most diverse on the planet

(Boyce et al., 2010). The Japanese Botanist, Mitsuru Hotta began fieldwork in Sarawak and

Brunei, concentrating especially on Schismatoglottis and its allies (Hotta, 1965). The studies

also continue within a few years by Josef Bogner and Niels Jacobsen which is still on going

work on the Schismatoglottideae and Cryptocoryne (Bogner, 1979, 1980; Jacobsen, 1985).

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Boyce et al. (2010) mentioned that several other botanists have entered the Borneo

aroid arena in the past 25 years. Alistar Hay started in the early 1980s by tackling the lasioids

as part of his doctoral research where as Peter Boyce in the mids 1980s. After that, he went on

to study and monograph Alocasia (Hay, 1998a, 2000), Colocasia (Hay, 1996),

Schismatoglottis (Hay & Yuzammi, 2000; Hay, 2002; Hay & Hersovitch, 2003) and

Homalomena (Hay & Hersovitch, 2002). Hiroshi Okada and Yasuko Mori which are another

two Japanese botanists that now working in Kalimantan, particularly on Schismatoglotideae

(Okada, 2000; Okada & Mori, 2000; Okada, Tsukaya & Mori, 1999). The first critical study of

evolution and taxagenesis in Sundaic aroids using a combination of extensive fieldwork and

molecular analysis was begun undertaking by Wong Sin Yeng. Besides, the generic

frameworks already changing according to her data and the understanding of the aroids of

Borneo also had forcing an examination.

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2.2 Tribe Schismatoglottideae

According to Hay & Yuzammi (2000), tribe Schismatoglottideae is a predominantly

Indomalayan taxon comprises 13 genera of terrestrial, lithophytic of rheophytic herbs with a

primary distribution centred on Borneo but with a total distribution ranging to SW Myanmar,

SW China, and Vanuatu. The largest genus, with over 200 species, is Schismatoglottis Zoll. &

Moritzi. All other genera are all much smaller : Apoballis Schott, Aridarum Ridl.,

Bucephalandra Schott, Bakoa P.C. Boyce & S.Y.Wong, Hestia S.Y.Wong & P.C. Boyce,

Hottarum Bogner & Nicolson, Ooia S.Y.Wong & P.C. Boyce, Phymatarum M.Hotta, Pichinia

S.Y.Wong & P.C. Boyce, Piptospatha N.E. Br., Schottariella P.C. Boyce & S.Y.Wong and

Schottarum P.C. Boyce & S.Y.Wong. Apart of these, nine genera are endemic on Borneo

which are Aridarum, Bucephalandra, Bakoa, Hottarum, Ooia, Phymatarum, Pichinia,

Schottariella and Schottarum. Most of Piptospatha species are mainly restricted to Borneo but

with two (non – Bornean) species occurring in Peninsular Malaysia and the far south of

Thailand. The single species of Hestia occurs in scattered localities along the west coast of

Peninsular Malaysia and also on Borneo. Apoballis, the next largest genus after

Schismatoglottis is predominantly Sumateran with a few species occurring on Jawa and in the

Peninsular Malaysia (Bogner & Hay, 2000; Boyce & Wong, 2008, 2009).

Schismatoglottis are commonly found on steep soil banks in forest where, perhaps, the

minute seeds can adhere and seedlings are less likely to be buried by leaf litter. Hay &

Yuzammi (2000) found a number of species are facultative (ex. S. tecturata) or obligate (ex. S.

multiflora) rheophytes, but no truly submerged aquatic species are known. Moreover, several

10

species are litophytic, mostly but not exclusively on limestones (ex. S. venusta, S. bauensis, S.

puberulipes). The seed testa of Schismatoglottideae usually costate and the embryo axile with

endosperm copious.

Recognition and description of Schismatoglottideae takes to three the number genera

that have been formally described for the Araceae tribes. They may be keyed out as follows:

1. Inflorescences erect to nodding at anthesis, if nodding, then either peduncle massive,

and peduncle at spathe insertion at most 45° from vertical axis. Infructescences fusiform with

a constricted orifice, if campanulate, then thick-walled and erect, never nodding. Plants of

various substrates but never on podsol …................………………………………………….. 2

– Inflorescences on very slender peduncles and nodding at anthesis. Peduncle at spathe

insertion flexing 180° from vertical axis. Infructescences narrowly campanulate, nodding.

Plants of pods …….………….…………………………………………..…...............…. Hestia

2. Modules monophyllous, congested in a distichous arrangement; ligular free sheath

persistent ….………………………………………………………………................... Pichinia

– Modules polyphyllous, never distichous; petiolar sheath fully attached, persistent or

marcescent …………………………………………...…………………......... Schismatoglottis

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For Figure 1 (a), Schismatoglottis jelandii species is most closely allied to Asperata

group which have the inflating of spathe and gaping of the limb when the female flowers are

active, then opening more or less wide as the male flowers shed their pollen. For Figure 1 (b),

species in Calyptrata group have the spathe limb falling just prior to the male flowers shedding

their pollen, abscising cleanly at the top of the lower spathe.

Figure 1: Inflorescences of genus of Schismatoglottis

(a): Inflorescence of Schismatoglottis jelandii at the early stage of male activity with wide-

open spathe limb (b): Spadix of Schismatoglottis calyptrata showing the female flower zone

partly fused to the spathe

(a) (b)

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2.3 Schismatoglottis

The aroids from family of Araceae are pre-eminent among the herbaceous flora of

Sarawak. Species of the genus Schismatoglottis are vegetatively similar with the genus of

Homalomena and the horticulturally popular genus Alocasia are among the most abundant and

diverse of the forest floor aroids. Schismatoglottis is a predominantly old world tropics genus

of more than 100 species of terrestrial herbaceous plants which also called as mesophytes,

which adapted to constant levels of soil and atmospheric moisture. However, some of this

genus is adapted to the flood zones of tropical forest streams (such plants are termed as

rheophytes). Others which very rarely grow as swamp plants in full sun called as helophytes.

The primary distribution of this genus is in Borneo.

The generic name of Schismatoglottis is from Greek Schisma, schismatos (separating)

and glotta (tongue) and refers to the variously deciduous upper part of the spathe of the most

species. Furthermore, the spathe in Schismatoglottis is differentiated into a lower persistent

portion enclosing the female zone of the spadix and an upper ephemeral spathe limb

subtending the male flower zone and terminal appendix. During the flowering time, the spathe

undergoes a series of movement, including inflating and spreading of the upper and lower

spathe, linked to managing pollinators during the various fertile phases of the flowers.

Besides, Schismatoglottis has a complex spadix structure and broadly divided into a lower

female flower zone, a middle male flower zone, which may or may not be separated from the

female zone by a sterile portion, and usually a terminal sterile appendix.

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The female flowers are often intermixed by irregularly arranged sterile male flowers.

In many species, these sterile structures are confined to a basal ring or to a single row along

the join between the spathe and the spadix. In other hand, the male flower zone of

Schismatoglottis consists of massed stamens not arranged or orientated into discrete male

flowers. The stamens are generally truncate and rose on short filaments or occasionally

stalkless with the filaments often but not always partially joined into small groups of two or

three. Schismatoglottis are divided into six informal taxonomic groups based on shoot

architecture, spathe structure, the structure of the petiolar sheath and the means by which the

spathe limb is shed. Four of these groups are present in Sarawak which is Calyptrata group,

Asperata group, Multiflora group and Tecturata group.

Hay (1996) has discussed the aspects of the morphology of Schismatoglottis in a

revision of the genus in the Malay Peninsula. However, some elaboration of the information is

required here as the revision which covers a much greater diversity of species. Moreover,

certain features have not been clearly found in some species due to lack of adequate materials.

Hotta (1965) has recognized five informal groups in the Bornean species, which are S.

homalomenoidea group, S. monoplacenta group, S. barbata group, S. acutifolia group and also

S. calyptrata group.

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The general key to informal species group may be keyed out as follows:

1. Stem hepaxanthic …………………………....………..……...…… S. calyptrata group

– Stem pleionanthic ………………………………………………………....................... 2

2. Spathe limb caducous ……………………………………………………………….... 3

- Spathe limb marcescent to crumbling and / or deliquescent ………………………….. 4

3. Leaf sheath fully attached to petiole …………………...…………. S. calyptrata group

- Leaf sheath ligular ……………………………………..……….… S. multiflora group

4. Petiole sheathing only at base; foliage leaves alternating with cataphylls

………………………………………………………………..........……..… S. tecturata group

- Petiole usually sheathing for at least a third of its length (rarely less); foliage leaves not

alternating with cataphylls ……………………………………………….............…………… 5

5. Spathe limb irregularly crumbling and breaking away at or after male anthesis

…………………………………………………………….................……… S. asperata group

- Spathe limb clasping the spadix and more or less marcescent after anthesis, finally

falling with spent parts of spadix ………………………………………............…………….. 6

6. Leaf sheath wings persistent; inflorescence nodding; male and female zones of spadix

more or less contiguous ………………………………………....................... S. corneri group

- Leaf sheath wings usually (but not always) deciduous; inflorescence more or less erect;

male and female zones separatedly nby conspicuous partly naked interstice

....…………………………………………………………...............……….. S. rupestris group

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2.4 Schismatoglottis calyptrata complex

According to Hay & Yuzammi (2000), Schismatoglottis calyptrata is the most

widespread species in the genus and a morphologically highly variable taxon. There are about

40 species under S. calyptrata complex. Based on the previous study, there are many species

published ranging from the minor variation in leaf blade shape to spadix morphology. Hay &

Yuzammi (2000) combined a considerable number of these names into much broadly-defined

species concept, although still recognizing that the existing localized population to appear to

be morphologically stable and indeed warrant recognition at a formal taxonomic rank. In

recent years an increase in fieldwork, especially in Borneo, has belief that at least some of the

published species names for S. calyptrata sensu lat. were too quickly classified (e.g., S.

muluensis M.Hotta) while at the same time that there exist in Sarawak a number of localized

populations that seemed to warrant taxonomis recognition.

Most of the Bornean species are divided between the Schismatoglottis calyptrata

group, the Schismatoglottis asperata group and the S. multiflora group. In addition, there are

three species with unclear affinity which are S. corneri, a gigantic arborescent pachycaul

species from northwestern Borneo with pleionanthic shoots, fully attached sheaths and huge

pendulous inflorescences with semi-persistent closed spathe limbs and apparently

unspecialised organization of the spadix. Next is S. tecturata, with very short leaf sheats and

foliage leaves alternating with cataphylls which take on the role of the leaf sheath and

persistent or marcescent spathe limbs. Lastly, S. petri which comes from Brunei may be

related to S. tecturata.

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The S. calyptrata group in Borneo has pleionanthic shoots including the remarkable S.

niahensis, which very rarely for this group. Besides S. calyptrata itself, there are three highly

variable widespread endemic species in Borneo, S. motleyana, S. trifasciata and S. trivittata

which are quiet difficult to distinguish from one another at their extreme variation. By

contrast, the rheophytic S. ahmadii, calcicolous S. viridissima, litophytic and calcicolous S.

venusta, and S. decipiens and S. silamensis confined to ultramafic substrate and others are also

some of localized species, several with specialized ecology.

Throughout the Old World range of the genus, Schismatoglottis calyptrata group have

39 species in Malaysia. S. calyptrata is the most widespread species in the genus but requires

ecological study throughout the range. This group usually is a hypogeal with 1-2 diameter and

hapaxanthic shoot with leaf sheath fully attached and persistent. Besides evergreen species,

sometimes S. calyptrata colony-forming small to massive and subarborescent herbs mostly of

rain forest floor or rheophytes or litophytes, with clear sap, 15-60 cm tall and the vegetative

tissue not or barely aromatic. Besides, S. calyptrata has 1-8 together inflorescences with a

strong acidic odour at female anthesis and the exposed part of 2-14 cm peduncle, erect at the

anthesis then deflected.

The blade is usually dull mid-green, sometimes variegated with 1-2 bands or

irregularly spotted grey-green to yellowish green; 7-35 cm long, widest at the base or ⅓ along

its length, and 4-18 cm wide, mostly cordate to sagittate. Sometimes it is oblong-lanceolate

with the base cordate to emerginate with posterior lobes rounded to rather sharply triangular

17

and 1-10cm long. However, the midrib abaxially prominent, with primary lateral veins 6-15

per side, irregularly alternating with lesser interprimaries, diverging at 45-70°, nearly always

raised adaxially towards the midrib, marginally impressed, entirely raised abaxially, mostly

not branched but occasionally with 1 or 2 branches especially in the lower part of the blade.

Meanwhile, the secondary venation of the S. calyptrata arising mostly from the midrib, but

sometimes from the bases of the primary veins and tertiary venation is inconspicuous.

A number of S. calyptrata has a 3.5-12 cm long spathe, lower spathe narrowly ovoid

with half of the length of the whole spathe green and often minutely longitudinally ridged and

asperous. The limb differentiated from lower spathe by an abrupt constriction corresponding

to the base of the male zone of the spadix, but at the female anthesis, the spathe much inflated,

narrowing and turbinate. The apex conspicuously mucronate, completely surrounding the

spadix and gaping ventrally or with the margins loosely overlapping with creamy to pale

greenish-yellow and also caducuous immediately after female anthesis.

The spadix comes from ¾ of the length of the spathe, narrowly hour-glassed shaped.

The female zone about half the length of the whole spadix, sessile, somewhat to markedly

obliquely inserted to partially adnate to spathe, 5-8 mm diameter below and distally tapering

to 3-4 mm diameter. Pistils are pale green, closely-packed with 1 mm tall and 0.5 mm

diameter, flask-shaped and close-packed below, distally becoming more widely spaced and

sunglobose finally rather widely scattered and squashed by the constricting spathe. The

stigmas look like a button-like, papillate, raised on a short style. Interpistillar staminodes are

white, mostly conspicuously taller than pistils, few in numbers, scattered, stalked, clavate,

rarely absent in very small forms.