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2015
CHARACTERIZATION OF NEST STRUCTURE,
THERMOREGULATION OF HIVE AND
MELISSOPALYNOLOGY OF GENIOTRIGONA
THORACICA SMITH (HYMENOPTERA: APIDAE;
MELIPONINI)
NUR FARISYA BINTI MOHD SAUFI
MASTER OF SCIENCE
2015
Characterization of Nest Structure, Thermoregulation of
Hive and Melissopalynology of Geniotrigona thoracica
Smith (Hymenoptera: Apidae; Meliponini)
by
Nur Farisya Binti Mohd Saufi
A thesis submitted in fulfillment of the requirements for the degree of
Master of Science
Faculty of Agro Based Industry
UNIVERSITI MALAYSIA KELANTAN
i
THESIS DECLARATION
I hereby certify that the work embodied in this thesis is the result of the original research and
has not been submitted for a higher degree to any other University or Institution.
OPEN ACCESS
EMBARGOES
CONFIDENTIAL
RESTRICTED
I agree that my thesis is to be made immediately available as
hardcopy or on-line open access (full text).
I agree that my thesis is to be made available as hardcopy or
on-line (full text) for a period approved by the Post Graduate
Committee.
Dated from until
(Contains confidential information under the office Official
Secret Act 1972)*
(Contains restricted information as specified by the
organization where research was done)*
I acknowledge that Universiti Malaysia Kelantan reserves the right as follows.
1. The thesis is the property of Universiti Malaysia Kelantan.
2. The library of Universiti Malaysia Kelantan has the right to make copies for the
purpose of research only.
3. The library has the right to make copies of the thesis for academic exchange.
SIGNATURE SIGNATURE OF SUPERVISOR
IC/ PASSPORT NO. NAME OF SUPERVISOR
Date: Date:
ii
ACKNOWLEDGEMENT
I am thankful to Dr. Kumara Thevan for having accepted to be my supervisor and for
providing me high quality scientific assistance. The pleasure and a great honor for me are
having you as my supervisor. You always motivated me throughout the study period and will
always remember the patience you had in supervising me in this research.
I am greatly indebted to Dr Arifullah for guidance in part of this research. The Faculty of
Agro Based Industry has provided the space and equipment that I needed for my lab work and
completes my thesis. This study received financial support from research grant R/RAGS/A
07.00/00760A/002/2012/000092.
I would like to give my appreciation to Prof Abu Hassan, Mr. Razip, Mdm. Wan Noriah for
their wise knowledge and suggestion about stingless bee. A big thanks to Mr Mohd Saupi
Pauzi and Mrs Yusnah Yusuf for sponsoring laptop in facilitates my research.
In additionally, I am sincerely would like to thank all my friends Husna, Vikram, Mr Ahmed,
Ilfah, Suaini, Sarah, Afiqah, Azwa, Aida, and Kelly for their support during my study. I will
cherish the friendship and memories.
Most importantly, I would like thank my beloved parents and siblings. Their support,
encouragement, patience and unconditionally love throughout my life and for allowing me to
further my study and carry out the research project smoothly.
iii
TABLE OF CONTENTS
PAGE
THESIS DECLARATION i
ACKNOWLEDGEMENT ii
TABLE OF CONTENTS iii
LIST OF TABLES vi
LIST OF FIGURES vii
LIST OF ABBREVIATIONS viii
LIST OF SYMBOLS ix
ABSTRAK x
ABSTRACT xi
CHAPTER 1 INTRODUCTION
1.1 Background of Study 1
1.2 Problem Statement 4
1.3 Significance of the Study 5
1.4 Objectives of the Study 6
CHAPTER 2 LITERATURE REVIEW
2.1 Stingless Bees 7
2.1.1
Origins and Historical Prospective
7
2.1.2
Distribution and Diversity of Stingless Bees
8
2.1.3
Life Cycle of Stingless Bees
9
2.1.4
Behaviour of Stingless Bees
10
2.2 Classification of Stingless Bee 12
2.2.1 Anatomy of Stingless Bee 14
2.3 Stingless Bees Nest 16
2.3.1 The Structure of Stingless Bees Nest 17
iv
2.3.2 Thermoregulation of Stingless Bees Nest 19
2.4 Economic Potential of Stingless Bees 20
2.4.1 Products of Stingless Bees 21
2.4.2 Pollination Service 23
CHAPTER 3 MATERIALS AND METHODS
3.1 Nest Characteristics of G. thoracica 26
3.1.1 Nest Structure and Nest Dynamics 26
3.1.2 Foraging Activity of Worker Bee 28
3.1.3 Morphometric Study of Worker Bee 29
3.2 Thermoregulation of G. thoracica 30
3.2.1 Thermoregulation of G. thoracica Hive 30
3.3 Melissopalynology of G. thoracica 31
3.3.1 Sample Collection 31
3.3.2
Acetolysis Procedure
31
3.4 Data Analysis 35 CHAPTER 4 RESULTS
4.1 Nest Characteristics of G. thoracica 36
4.1.1 Nest Structure of G. thoracica 36
4.1.2 Nest Dynamics of G. thoracica 40
4.1.3 Foraging Activity of G. thoracica 42
4.1.4 Morphometric Parameters of G. thoracica 45
4.1.5 Flight Cost and Aggressive Cost of G. thoracica 47
4.2 Thermoregulation of G. thoracica 48
4.2.1 Monthly Temperature and Monthly Humidity 48
4.2.2 Comparison between Ambient Condition and Hive Condition 50
4.3 Melissopalynology of G. thoracica 51
v
4.3.1 Analysis of Pollen in G. thoracica Honey 51
4.3.2 Types of Honey Produced by G. thoracica 58
4.3.3 Pollen Calendar of G. thoracica Honey (Agropark UMK) 63
CHAPTER 5 DISCUSSION
5.1 Nest Characteristics of G. thoracica 66
5.2 Thermoregulation of G. thoracica 70
5.3 Melissopalynology of G. thoracica 72
CHAPTER 6 CONCLUSION AND FUTURE WORK
6.1 Research Findings 75
6.2 Conclusion 76
6.3 Recommendation 78
REFERENCES 79
APPENDIX A 86
APPENDIX B 88
APPENDIX C 91
APPENDIX D 95
APPENDIX E 99
LIST OF PUBLICATIONS 100
vi
LIST OF TABLES
NO. PAGE
4.1 Size of external and internal nest structure of G. thoracica 41
4.2 Analysis of foraging activity of foragers 49
4.3 Measurement of morphometry parameters of G. thoracica 50
4.4 Monthly mean temperature and mean relative humidity for 54
thermoregulation of G. thoracica
4.5 List of pollen types found in honey of G. thoracica 57
4.6 Unifloral honey produced by G. thoracica by month based on pollen 65
frequency
4.7 Multifloral honey with secondary pollen types produced by G. thoracica 66
by month based on pollen frequency
4.8 Multifloral honey produced by G. thoracica by month based on pollen 68
frequency
4.9 Pollen calendar of G. thoracica honey 2013/2014 70
vii
LIST OF FIGURES
NO. PAGE
2.2 Anatomy of stingless bee 16
2.3 General nest structure of stingless bee in tree trunk 21
3.1 Design of artificial hive of G. thoracica. 30
3.2 Flow chart of Image J analysis for nest dynamics of G. thoracica 31
3.3 Flow chart of acetolysis for melissopalynology of G. thoracica 37
4.1 External and internal nest structure of G. thoracica 42
4.2 The illustration of top view of internal nest structure of G. thoracica 43
4.3 Internal nest structure of G. thoracica with invulcrum or batumen 44
4.5 Nest dynamics of internal nest components (brood combs area, honey pots 46
and pollen pots area) of G. thoracica by month
4.6 Monthly honey volume from three colonies of G. thoracica 47
4.7 Foraging activities of G. thoracica foragers 48
4.8 Anatomy parts used in morphometry study of G. thoracica worker 51
4.9 Pollen types from non-seasonal tree in G. thoracica honey 58
4.10 Pollen types from seasonal tree in G. thoracica honey 59
4.11 Pollen types that from shrub in G. thoracica honey 60
4.12 Pollen types from herbs in G. thoracica honey 61
4.13 Pollen types from vegetables in G. thoracica honey 62
4.14 Pollen types from grass in G. thoracica honey 62
viii
LIST OF ABBREVIATIONS
T. Trigona
G. Geniotrigona
M. Melipona
mm millimeter
USD United States Dollar
w width
h height
hrs. hours
l length
g gram
mg milligram
ml milliliter
µm micrometer
d diameter
A. Apis
N North
E East
No. number
SD standard deviation
ix
LIST OF SYMBOLS
ºC degree Celsius
% percentage
(=) synonymous names
x
Ciri-ciri Struktur Sarang, Termoregulasi Sarang dan Melisopalinologi Geniotrigona
thoracica Smith (Hymenoptera: Apidae; Meliponini)
ABSTRAK
Kajian mengenai struktur sarang lebah kelulut bagi spesis yang terpilih dapat memberi
manfaat kepada industri kultur-meliponi. Kultur-meliponi adalah aktiviti menternak lebah
kelulut. Genotrigona thoracica merupakan salah satu lebah kelulut yang bersaiz besar di
Malaysia dan mempunyai nilai ekonomi dalam kultur-meliponi. Objektif kajian ini adalah
untuk mengenalpasti struktur sarang dan dinamik sarang, aktiviti keluar masuk dan masa
kekerapan keluar masuk dan morphometri lebah pekerja; mengenalpasti suhu sarang dan
analisis kebolehan termoregulasi; mengenalpasti jenis debunga di dalam madu, jenis madu
yang dihasilkan dan kalendar debunga oleh G. thoracica. Koloni (n=3) G. thoracica
digunakan dalam mengkaji struktur sarang dan dinamik sarang, aktiviti lebah kelulut keluar
dari sarang dan kajian morphometri lebah pekerja, thermoregulasi dan melisopalinologi, iaitu
analisis debunga di dalam madu. Persempelan telah dilakukan dari Julai 2013 sehingga Jun
2014 untuk kajian dinamik sarang serta aktiviti lebah kelulut keluar dari sarang,
thermoregulasi dan melisopalinologi. Untuk kajian morphometri, sampel lebah pekerja telah
diambil dari setiap sarang selama lima bulan bermula dari Februari 2013 sehingga Jun 2014.
Hasil kajian menunjukkan struktur luaran sarang terbentuk seperti timbunan berwarna perang
dan struktur dalaman sarang terdiri dari tiga komponen iaitu susunan telur yang dikelilingi
oleh kantung madu dan debunga. Dinamik sarang G. thoracica berubah mengikut musim.
Selain itu, waktu puncak bagi kekerapan G. thoracica keluar dari sarang adalah dalam
tempoh 1000 jam sehingga 1200 jam. Dari kajian morphometri lebah pekerja, menunjukkan
terdapat perbezaan yang signifikan, p < 0.05 untuk bahagian anatomi dalam tempoh lima
bulan persempelan. Kajian morphometri ini juga menunjukkan G. thoracica mempunyai kos
penerbangan yang tinggi berbanding kos agressif. Keputusan termoregulasi menunjukkan G.
thoracica tidak mempunyai keupayaan untuk termoregulasi dimana tiada perbezaan
signifikan, p > 0.05 antara suhu sarang dengan suhu persekitaran. Selain itu, kajian
melisopalinologi menunjukkan terdapat 30 debunga yang terdapat dalam sampel madu G.
thoracica dan tiada perbezaan signifikan p > 0.05 antara debunga yang terdapat di dalam
madu sepanjang tahun. Tambahan, dari hasil kajian melisopalinologi menunjukkan dua jenis
madu yang dihasilkan oleh G. thoracica iaitu uniflora dan multiflora pada bulan yang berbeza
sepanjang tahun. Kesimpulannya, ciri-ciri struktur sarang, kebolehan thermoregulasi dan
melissopalinologi bagi G. thoracica telah diperolehi.
xi
Characterization of Nest Structure, Thermoregulation of Hive and Melissopalynology of
Geniotrigona thoracica Smith (Hymenoptera: Apidae; Meliponini)
ABSTRACT
The study on stingless bee nest structure of selected species brings benefit to meliponiculture.
Meliponiculture is an activity of stingless bee keeping. Geniotrigona thoracica, is one of the
largest stingless bee in Malaysia and has economic potential used in meliponiculture. The
objectives of this study are to determine nest structure and nest dynamics, foraging activity
and morphometry of worker bee; determine hive temperature and analyses thermoregulation
ability; identify pollen types in honey, types of honey produced and pollen calendar of G.
thoracica. Colonies (n=3) of G. thoracica were used in study of nest structure and nest
dynamics, foraging activity and morphometry study of worker bee, thermoregulation and
melissopalynology, analysis of pollen in honey. Sampling was done from July 2013 until
June 2014 for nest dynamics and foraging activity, thermoregulation and melissopalynology.
For morphometry study, worker bees were sampled from each nest for five months, from
February 2014 until June 2014. Result showed external nest structure formed in brown mount
shape and the internal nest formed by three components, horizontal brood combs that
surround with honey pots and pollen pots. The nest dynamics of G. thoracica changes
according to the season. The peak time foraging activity of G. thoracica was in between 1000
hrs until 1200 hrs. From morphometry study of worker bee, showed there was significant
difference, p < 0.05 of anatomy parts in five months of sampling. This morphometry study
also showed that G. thoracica having high flight cost than aggressive cost. The result showed
that G. thoracica does not have ability to thermoregulate, where there was no significant
difference p > 0.05 between hive and ambient temperature. Then, the melissopalynology
study showed that there were 30 pollen grains were present in sample of G. thoracica honey
and there was no significant difference, p > 0.05 between types of pollen present in honey
throughout a year. Furthermore, from this melissopalynology study, two types of honey
produced by G. thoracica were unifloral and multifloral in different month throughout a year.
In conclusion, nest characterization, thermoregulation ability and melissopalynology of G.
thoracica were determined.
1
CHAPTER 1
INTRODUCTION
1.1 Background of Study
Stingless bees are social bees which lack a functional sting and belong to the
tribe Meliponini (Michener, 2013). They are able to produce honey similar to honey
bees. In Malaysia there were recorded about 16 to 32 identified species of stingless
bees (Eltz & Bru, 2003; Hannah et al., 2012; Norowi et al., 2010). The
meliponiculture is an activity of beekeeping with stingless bees by which bee keepers
maintain, propagate and utilize stingless bee colonies of various species for profit
(Halcroft et al., 2013a). The stingless bees colonies managed in artificial hives
enables bee keepers to propagate colonies and also to produce products such as honey,
pollen, cerumen and propolis (Vijayakumar et al., 2013). The meliponiculture
industry is new in Malaysia, whereas this activity widely practiced in Brazil, Mexico,
Africa, Australia and Thailand (Cortopassi-Laurino et al., 2006; Halcroft et al., 2013c;
Sawatthum, 2004). Meliponiculture allow bee farmers to generate income by selling
stingless bee colonies, honey, bee bread, propolis, pollination services, educational
services and agro-tourism (Halcroft et al., 2013d; Steve, 2013).
Though there were many stingless bee species in Malaysia, not all species
have economic value. In Australia, there are two species of stingless dominated in
meliponiculture industry which are Tetragonula carbonaria and Austroplebeia
australis (Halcroft et al., 2013a). Meanwhile, in Thailand there are four species of
2
economic value, which are Trigona pegdeni Schwarz, Tetragonula laeviceps,
Lepidotrigona terminata Smith and Lepidotrigona ventralis doipaensis Schwarz,
where these species of stingless bees used for pollination services (Sawatthum, 2004).
Recent study showed that, there are five species of stingless bees have commercial
value in Malaysia. They are Geniotrigona thoracica Smith, Heterotrigona itama
Cockerell, L. terminata Smith, Tetragonula fuscobalteata Cameron and T. laeviceps
(Kelly et al., 2014). In order to sustain the meliponiculture industry in Malaysia,
research on their nest biology and behaviour of stingless bees were needed. There still
lack of information regarding the nest structure of economic potential of stingless bee
species in Malaysia. Thus, one of the objectives of this research is to determine the
nest structure of one of economic potential stingless bees, Geniotrigona thoracica.
Since, nest stingless bees mainly found in tree hollows, it is difficult for bee
farmers to extract nest products. Hence, by transferring the colony of stingless bees
into the artificial hive ease the process of extracting nest product. Moreover, it is
easier to move and propagate the stingless bees colonies (Cortopassi-Laurino et al.,
2006). There are a few designs for artificial hive in meliponiculture. For example, in
India, traditional beekeeping of stingless bee by Kani tribes use a bamboo pole
beehive to keep the stingless bee of Tetragonula irridipennis (Kumar et al., 2012).
Others, design for artificial hives called „Utrecht University-Tobago Hive‟ (UTOB)
design, where the hive separate the honey section from the brood section (Sommet,
1999). There are few factors to be taken into consideration when constructing
stingless beehive. These factors are; the volume of the natural nest, orientation of hive
(vertical or horizontal) and position of nest entrance which varies by species
(Kwapong et al., 2010). It is hypothesized in this research that these designs of
3
artificial hive play important role for adaptation of stingless bee especially in
thermoregulation process. Thermoregulation is found among large-bodied insects
where has significant ecological consequences because the internal heat generation
enables solitary and social bees to forage and pollinate under colder surrounding
temperature (Nieh & Sánchez, 2005). It is important for bees maintain the thermal
condition of their nest in order to promote the successful breeding in their colonies.
Researchers found only few species of stingless bees are able to maintain the
temperature in the nest (Sung et al., 2008). Thus, one of the aims of this research was
to determine the ability of G. thoracica to thermoregulate inside the artificial hive.
In Malaysia, the demand for stingless bee honey has increased and being the
commercial value higher than the one of Apis melifera honey. Similar situation have
been documented in Thailand (Seanbualang, 2012). Honey production of stingless
bees depends on the species of stingless bee, the available of resources and weather.
Microscopic analysis of pollen grains of natural honeys known as melissopalynology,
which allows identification of the different nectar sources over the season. Bees use
these sources for the production of honey in a region, classifying the honey
botanically and geographically according to its origin (Upadhyay et al., 2014).
Through the identification of pollen occur in stingless bees honey sample allow the
characterization of vegetative surround the stingless bees nest and at the same time
contribute to pollination (Louveaux et al., 1970). In addition, bees are the most
important insect pollinator because a wide variety of them are known to be efficient
and effective pollinators of many plant species, but if the later honey bee is declining
as been reported, stingless bees could be important pollinator to replace honey bee
(Biesmeijer et al., 2006; FAO, 2007). Besides, a research on utilization of stingless
4
bees for pollination services in agricultural ecosystems in Malaysia showed, four
stingless bees species, Heterotrigona itama, Geniotrigona thoracica, Tetragonula
atripes and Tetrigona peninsularis can be potentially domesticated for pollination
service (Norowi et al., 2010). Hence, pollen analysis is important step in identifying
plant that preferred by stingless bees in honey production (Louveaux et al., 1970). In
this research, monthly honey sample of G. thoracica harvested and pollen occur in
honey were analysed in order to determine types of pollen present and characterize the
sample of honey produced.
Meliponiculture industry in Malaysia has booming potential. Research on
selected stingless bees species which is suitable used to domesticate in this industry
will help beekeepers to maintain and develop the meliponiculture industry. Thus, this
research will focus on one of the largest stingless bee species in Malaysia, G.
thoracica, and determine their nest structure, thermoregulation of the nest and
melissopalynology.
1.2 Problem Statement
This research mainly focuses on one of the largest stingless bee species in
Malaysia, G. thoracica. In meliponiculture industry, bee farmers were having lack of
information or knowledge in biology of this species. Result from this research will
provide information on the nest structure of this stingless bee species which will help
bee farmers to know about nature of this species.
5
The artificial hive is one the main element use in meliponiculture, design and
material for construction of artificial hive are important to maintain and propagate the
stingless bee colony (Cortopassi-Laurino et al., 2006). The problem will arise when
the stingless bee colonies are being transferred into inappropriate artificial hive. The
design and materials for construction of artificial hive need to take consideration of
ability of this stingless bee to thermoregulate. Hence, research on design of artificial
hive and ability of stingless bee to thermoregulate are needed in maintain the
meliponiculture industry.
The availability of food source around the bee farm is important to maintain
the stingless bee colonies. The problem in meliponiculture activity is depletion of
food source for stingless bee especially during raining season. Study on pollen
collected or pollen occurrence in honey sample may overcome the problem of
depletion of food source and also to maintain ecosystem of the bee farms. In addition,
research on pollen favour by stingless bee may help in pollination service.
Therefore, research on characterization of nest, thermoregulation of hive and
melissopalynology of stingless bees are needed in solve some of the problems arise in
meliponiculture industry in Malaysia.
1.3 Significance of the Study
This research contributes to the knowledge on the economic potential of
stingless bee species in Malaysia, G. thoracica. The result of this research helps bee
farmers to understand their nest structure. The study of internal and external nest
structure will give idea on how G. thoracica arrange their internal nest component.
6
This information will be useful in transferring the colony of G. thoracica into artificial
hive.
Besides, monthly data of nest dynamics of G. thoracica will give idea on the
survival rate of G. thoracica inside artificial hive. These data also will indicate which
product mostly produces by this species; honey, propolis or bee bread (= pollen).
Meanwhile, thermoregulation of G. thoracica indicate that this species of
stingless bees able to adapt with the micro-climate at the site of research. The result
from thermoregulation from this research give some information about design of the
artificial hive used in this research is suitable for G. thoracica or not.
Melissopalynolgy or pollen analysis using honey sample will give idea on
favoured stingless bee plant in honey production. Result from this research will
contribute the knowledge on plant that favoured by G. thoracica in honey production.
1.4 Objectives of the Study
1. To determine nest structure and nest dynamics, foraging activity and identify
morphometry of worker bee of G. thoracica.
2. To determine the hive temperature and analyse the thermoregulation ability of G.
thoracica.
3. To identify the pollen type in honey of G. thoracica and to determine the types of
honey produced by G. thoracica also to construct pollen calendar.
7
CHAPTER 2
LITERATURE REVIEW
2.1 Stingless Bees
Stingless bees are social bees that belong to the order Hymenoptera and one of
four tribes (Meliponini) in the family Apidae. Term “stingless bees” referred to all
male bees are completely stingless, the sting is reduced and modified ovipositor and
do not function for stinging that only found in female bees (Michener, 2013).
Although stingless bee do not sting, but they will defend themselves by biting when
their nest is disturbed (Wille, 1983). In addition, a few in the genus Oxytrigona and
several other species of the genus Trigona have mandibular secretions that cause long
lasting lesions (Michener, 2013).
2.1.1 Origins and Historical Prospective
Stingless bees are considered originated in South America where can be found
great diversity of stingless bees group and meliponiculture is extensively practiced
there (Eardley, 2004). Africa has been suggested to be the centre of stingless bees‟
origin which then has dispersed to other tropical and subtropical parts of the world
and this suggestion is based on paleontological and biogeographical data and also
supported by the fact that their primitive species with a well-developed sting system
8
are found in Africa (Wille, 1983). Before the discovery and the conquest of the
American continent by Europeans, the rearing of stingless bees was an important part
of the commercial and alimentary customs of many indigenous cultures of Central and
South America (Cortopassi-Laurino et al., 2006).
In view of the growing pressure on the environment and the associated loss of
honeybees, attention has focused on meliponiculture as an eco-friendly agro-based
venture (Slaa & Sanchez, 2006). Stingless bees are known to be one of the key
contributors in providing ecosystem support services vital to the survival of several
forest plant and crop species through pollination (Biesmeijer et al., 2006; FAO, 2007;
Slaa & Sanchez, 2006).
2.1.2 Distribution and Diversity of Stingless Bees
Stingless bees are found throughout most of the tropical and subtropical
regions of the world such as Australia, Africa, Southeast Asia, and in some parts of
South America (Mexico, Brazil, Peru, Venezuela, Paraguay, Bolivia and Costa Rica)
(Maria et al., 2013). Approximate numbers of species so far identified were 50 species
in Africa, 300 species in the Americas, 60 species in Asia, ten species in Australia and
four species in Madagascar (Bradbear, 2009). In Asia there were 43 recognized
species belonging to two genera which were Lisotrigona and Trigona (Michener,
2007). The genus Trigona comprising about 120 species and were placed into ten
subgenera which were Homotrigona, Lepidotrigona and Heterotrigona (Chinh et al.,
2005). Malaysia hosts wide variety of stingless bees, there were about 32 species of
stingless bees were identified (Norowi et al., 2010). The studies on distributions of
9
stingless bees in Malaysia were located according to study locations. These locations
were Sabah, Negeri Sembilan, Selangor, Perak and Terengganu (Eltz & Bru, 2003;
Hannah et al., 2012).
2.1.3 Life Cycle of Stingless Bees
There is no solitary phase in meliponine life cycle, where colony life is
continuous. When a colony is dividing, workers from parent colony fly to a new site
and prepare it as a nest. These workers fly in repeated trips carrying construction
materials and food to new nest (Michener, 2013). An unmated young queen (= gynes)
flies to new nest from parent colony. The interchange process took a few months until
the new colony become independent. During this process, many males (= drones)
often from other colonies assemble near or hover near the nest entrance presume that
attracted by pheromones produced by young queen. Meliponine queens normally mate
only once, after mating a gyne may become a queen either replace the old queen or
serves in new colony (Chinh, 2004).
In Meliponini, brood production involves a sequence of interactions between
the queen and a group of workers. This process termed the “provisioning and
ovipositing process” or “POP”. Brood cells were mass-provisioned with a mixture of
honey, pollen, and protein rich secretions from the hypopharyngeal glands. Once the
cells were provisioned, the queen oviposits and workers sealed the cell. Some species
provision cells successively while others provision synchronously (Halcroft et al.,
2013d).
10
A diploid female is produced when a sperm cell is released to fertilize the egg
as it passes through the oviduct (Halcroft et al., 2013b). Research showed that the
production of male are depending on seasons and availability of food stored in colony
(Chinh, 2004). Besides, the amount mass provisioned into cells also will determine
the caste. Queens are nurtured in larger brood cells, called royal cells, whereas males
and workers are nurtured in smaller brood cells. Generally, royal cells are provisioned
with up to eight times more larval food than worker and male cell (Menezes et al.,
2013). Thus, female larvae will develop into queens if they receive the large amount
of larval food.
Adults stingless bees emerge as workers are performed most of the activities
of the nest such as storage pot construction, honey and pollen processing and packing,
house cleaning and maintenance, larval feeding and provisioning, queen grooming
and feeding, and colony security. Meanwhile, queen and drone only involve in mating
(Kwapong et al., 2010). There is limited study on life span of stingless bees, studies
reported that life span of worker bee in the range of 60-161 days (Halcroft et al.,
2013a).
2.1.4 Behaviour of Stingless Bees
Foraging is a part of colony behaviour of stingless bees (Kwapong et al.,
2010). During foraging, stingless bees collect pollen, nectar, oils, water, resins, muds
and sand particles. Most stingless bees species will communicate with each other on
locations of forage sources by secreting pheromones (Michener, 2013). The
pheromones used by several species of stingless bees for scent trail communication
are generally assumed to be produced by mandibular glands. A study showed that in
11
Trigona recursa these pheromones originate from the labial glands, which are well
developed in the heads of foragers (Jarau et al., 2010). When the foraging workers
encounter any resources, they will collect and return to their nest to recruit other
workers to the forage source. During the trip, workers stop and mark specific spots
with pheromones in order to direct the other workers to the forage source (Sánchez &
Vandame, 2013). Workers bees begin foraging activities as early as dawn and end by
dusk depending on weather conditions and availability of forage. Peak foraging times
happen simultaneously on dry season when the forage sources is abundance
(Kwapong et al., 2010). Meanwhile, research on T. irridipennis showed the foraging
activity is higher during winter and the forager that return to nest with pollen load
higher during summer (Danaraddi, 2007).
Guarding behaviour also part of colony behaviour of stingless bees (Kwapong
et al., 2010). Generally, guards occur in small numbers within the entrance of the nest
but they withdraw into the entrance tube if too closely observed. Guards are not
normally aggressive towards human observers. However, if the nest is opened
workers can become moderately to strongly aggressive. They buzz around the heads
of human “predators” and spot their hair with globules of resin until the nest is sealed
(Halcroft et al., 2013c). Research showed the number of standing guards more than
hovering guards. This might simply be because standing on the entrance tube is
energetically more affordable for colonies than hovering. In Tetragonisca angustula,
breath did not cause aggressive defensive reactions. Instead, the number of guarding
bees rapidly decreased as the standing guards retreated into the nest (Zweden et al.,
2011). In the night, workers on guard retreat into the nest and use propolis to close
their nest entrances. This behaviour prevents ants, beetles and other animals from
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entering the nest. The propolis seal at the entrance is removed in the morning and new
ones replaced every evening (Kwapong et al., 2010). Some species of stingless bees
use sticky plant materials such as resins to prevent intruders from entering their nests.
Where intruders or predators manage to enter nests, large volumes of resins and
propolis are used to entomb them alive till they die (Zweden et al., 2011).
2.2 Classification of Stingless Bee
The stingless bees are classified into 23 genera and 18 subgenera, it was
estimated that 400 to 500 different species of stingless bees were known, but new
species were identified every year (Michener, 2007). Earlier researchers placed all
Meliponini in a single genus, Melipona (Michener, 2013). Then, others recognized
two major genera, Melipona for the species now placed in that genus and Trigona for
all the rest of the Meliponini except a few robber species commonly placed in a
separate genus. Several groups were named as subgenera of Trigona but in 1946 and
thereafter Moure elevated subgeneric groups to the status of genera and described
various new genera (Michener, 2013). At a moment, the meliponines were discussed
as subfamily Meliponinae, with tribe Meliponini Lepetier, Lestrimelittini Moure and
Trigonini Moure. After 1990, the Meliponini were considered by some researchers to
be a single tribe containing all stingless honey making bees, in a one subfamily,
Apinae, containing these and the bumble bees, honey bees and the euglossine
(Roubik, 2013).
Identification of stingless bees to the genus and subgenus levels should be
facilitated by the keys of Moure (1961) and (Michener, 2000, 2007) then,
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identification of stingless bees to the species level is made possible by revisional
works such (Schwarz, 1937, 1939) and (Sakagami, 1975, 1978) for Asian region
(Michener, 2013).
Recent research showed, the valid names for stingless bees are now 32 genera
in the Neotropics, 15 in Asian and Australian region and nine in Afrotropics and a
total including all names and fossil meliponines of 61 genera. When, more fossils
being more discovered, a few more living genera that is likely to change. Nowadays,
the stingless bees number approximately 56 genera (Roubik, 2013). Based on
molecular study of stingless bee phylogeny, there are 165 different taxa from 55 of 61
world genera which divided into three clades: Afrotropical, Indo-Malay/ Australasian
and Neotropic. The genera lies on Neotropic clade are Paratrigona, Aparatrigona,
Nogueirapis, Partamona, Parapartamona, Cephalotrigona, Trigona, Ptilotrigona,
Tetragona, Geotrigona, Oxytrigona, Scaptotrigona, Duckeola, Trichotrigona,
Frieseomelitta, Tetragonisca, Friesella, Plebeia, Lestrimelitta, Plebeia, Mourella,
Schwarziana, Nannotrigona, Scaura, Schwarzula, Eomelipona, Michmelia,
Melikerria, Melipona, Eomelipona, Celetrigona, Dolichotrigona, Trigonisca, and
Leurotrigona. Genera that can be found in Indo- Malay/ Australasian are
Geniotrigona, Lepidotrigona, Heterotrigona, Platytrigona, Sundatrigona,
Homotrigona, Tetrigona, Odontotrigona, Lophotrigona, Tetragonilla ,Tetragonula,
Lisotrigona, and Austroplebeia. Genera Hypotrigona, Liotrigona, Lisotrigona,
Austroplebeia, Apotrigona, Meliplebeia, Axestotrigona, Plebeiella, Dactylurina,
Meliponula and Plebeina lies on clade Aftrotropical (Rasmussen & Cameron, 2009).
