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The Trigona species (stingless bees), their important and role of pollination in the higher plants Arbind Mani Tripathi [email protected] Horticulture Department Mejo University, Thailand

Introduction Bees are social insects which are closely related to wasps and ants. The bodies of bees are divided into head, thorax and abdomen, with three pairs of legs, pairs of segmented antennae and two pairs of wings on the thorax. The queen lays eggs in the wax cell by the ovipositor that is modified in the workers to form a sting. Bees are living in colonies. They have division of labour in which individuals carry out particular duties. There are three kinds of bee in a colony, a few hundred drones or males, one or two egg-laying female or queen, and from 300 to 100,000 thousand workers (bees’ number are varied on species and environment) [Anonymous, 2009e, Baconawa, 2009]. The mature queen is easily recognized by her large abdomen. Bees have  a   combined   mouth   parts,  so  they   can   both   chew   and   suck  (whereas grasshoppers can chew and moth can suck, but not both) (Anonymous 2009c).  Honey bees are primarily distinguished by the producing honey, beeswax and pollination. Honey bees are belongs to the members of the tribe Apini and genus Apis. They represent only a small part of the approximately 20,000 known species of bees (Anonymous, 2009a, Anonymous, 2009b). Also some other types of related bees produce and store honey, but only members of the genus Apis are true honey bees. The largest bee in the world is a Indonesian resin bee or leafcutter bee (Megachile plutowhose) which females can attain a length of 39 mm while the smallest bee is Trigona minima, a stingless bee those workers are about 2.1 mm long (Anonymous, 2009a). Stingless bees Stingless bees are very diverse in behaviour, but they are highly eusocial. Stingless bees are the tribe of Meliponini in the family Apidae, and closely related to the common honey bees and found in most tropical or subtropical regions of the world. Stingless bees are not active all year round; they are less active in cooler weather. Unlike other eusocial bees, they do not sting but will defend by biting if their nest is disturbed. They are living usually nest in hollow trunks, tree branches, underground cavities, or rock crevices. 500 stingless bees’ species are recorded and they are classified into five genera: Melipona, Trigona, Meliponula, Dectylurina and Lestrimelitta and some of them like Trigona spp. and Melipona spp. are the honey producing bees. Australian stingless (Teragonula carbonaria) bees produce less than one kilogram honey but that is the prized as a medicine in many African communities (Anonymous, 2009e). Trigona species Trigona is the largest genus of stingless bees and have many subgenera. Trigona is a genus of the Meliponini tribe which is found extensively in tropical regions. It extends from Mexico to Argentina, India, Sri Lanka to Taiwan, the Solomon Islands, South Indonesia and New Guinea (Michener, 2000 cited by Klakasikorn et al. 2005) but no members of the genus occur in Africa (Anonymous, 2009f). The two species(Trigona binghami and Trigona minor) are new added to the list of 30 species recorded earlier by Schwarz (1939), Sakagami et al. (1985) and Michener and Boongird (2004) cited by Klakasikorn et al. (2005) making a total of 32 stingless bees in Trigona species are currently recorded from Thailand. The newly

recorded species were found in HM Queen Sirikit Botanical Garden in Maerim, Chiang Mai, Chanthaburi and Mae Hong Son Provinces, Thailand (Klakasikorn et al. 2005). During 2004 to March 2005 in the lower mixed deciduous forest at the Golden Jubilee Thong Pha Phum Project, in Kanchanaburi Province, 2 genera (Trigona spp. and Hypotrigona spp.) and sixteen species of the stingless bees were found in this area, namely Trigona apicalis Smith, T. melanoleuca Cockerell, T. atripes Smith, T. canifrons Smith, T. thoracica Smith, T. terminata Smith, T. ventralis Smith, T. flavibasis Cockerell, T. iridipennis variety 1, T. iridipennis variety 2, T. iridipennis variety 3, T. iridipennis variety 4, Hypotrigona scintillans, H. pendleburyi and H. klossi . These Trigona species’ morphological characters are given in fig. 1. The diversity of Trigona spp. and their resin and gum collecting behaviour mostly depended on environmental factors. The bees preferred to collect resin and gum from 16 plant families including Anacardiceae, Dipterocarpaceae, Euphobiaceae, Hypericaceae, Meliaceae and Moraceae. During the rainy season they collected resin and gum all day time while during the dry season start to afternoon until late in the day. T. apicalis collected resin and gum to make the largest number of propolis compared with the other bee species (Inson and Malaipan, 2006). Fig.

( 1 )

Trigona apicalis Smith (2) T. melanoleuca Cockerell (3) T. peninsularis Cockerell (4) T. collina Smith (5) T. canifrons Smith (6) T. thoracica Smith (7) T. terminate Smith (8) T. ventralis Smith (9) T. flavibasis Cockerell (10) T. iridipennis variety 1 (11) T. iridipennis variety 2 (12) T. iridipennis variety 3 (13) T. iridipennis variety 4 (14) Hypotrigona scintillans (15) H. pendleburyi variety 1(16) H. klossi

The cytogenetic study of 31 species of genus Trigona by the Costa et al. 2004 and found (all of the species) females had 2n = 34chromosomes and males had n = 17 chromosomes. The C-banding patterns showed that the karyotypes of these species consisted mainly of acrocentric and pseudoacrocentric chromosomes. Pollination Pollination means the transfer of pollen grains from anther to stigma by the help of biotic and abiotic agents with same plant or same kind of species. After reaching of pollen grains in plants then fertilization or sexual reproduction take place. Pollination is a necessary step in the reproduction of flowering plants and resulting in the production of offspring. The pollination process as an interaction between flower and vector was first addressed in the 18th century by Christian Konrad Sprengel (Anonymous, 2009d). It is important in horticultural crops because of fruits are the end product of pollination which gets after fertilization. There are two types of pollination such as abiotic and biotic pollination. Abiotic pollination Abiotic pollination means where pollination is taking place without the involvement of living organisms or transfer of pollen by help of wind and water. It’s quite rare, only 10% of flowering plants are pollinated without organisms. This form of pollination is predominant in grasses, most conifers, and many deciduous trees. In the aquatic plants pollination being through water which releases their pollen directly into the surrounding water. About 80% of all plant pollination is biotic. Of the 20% of abiotically pollinated species, 98% is by wind and 2% by water (Anonymous, 2009d). Biotic pollination It is antonyms of abiotic pollination or it means when pollination is taking place by the help of living organisms like insect, birds, etc. are called biotic pollination. There are roughly 200,000 varieties of animal pollinators in the wild; most of them are insects (Anonymous, 2009d). Pollination by insects (bees, wasps, ants, beetles, moths and butterflies and flies) often occurs on plants due to their attractive developed colour petals and a strong scent. And also pollination is doing by vertebrates such as birds and bats. Plants adapted to using bats or moths as pollinators typically have white petals and a strong scent. Mechanics of pollination Some times the terms "pollinator" and "pollenizer" are confused, for clear understanding the term pollinator means the agent of biotic or abiotic things which are help to transfer the pollen from source to definite place while term pollenizer is the plant that develops as the pollen source for same or other plants. Some plants are self-compatible and can pollinate themselves while other plants have chemical or physical barriers to selfpollination and need to be cross-pollination. In pollination management, a good pollenizer is a plant that provides compatible, viable and plentiful pollen and blooms at the same time because of pollination requires consideration of pollenizers. According to pollenizers, pollination can be classified in two ways like cross-pollination (with a pollinator and an external pollenizer) and self-pollination (basically without pollinator). Plants and pollinator interaction due to biological and physical features such as colour shape and odour of the flower and it is governed by the energy needs. The pollinators are highly selective in their floral visits and shown to choose those flowers which best meet their energetic needs. The energy needs and foraging dynamics of pollinators are dependent upon prevailing weather conditions which regulate the schedule of pollination activities (Abrol, 2005). Pollinators and agriculture

Pollinators provide an important ecosystem service to both natural and agricultural ecosystems. Pollinators ensure fruit set development and dispersal in the vast majority plants in both ecosystems. In turn, plants provide food and nesting resources for pollinators. The process of securing effective pollinators to service agricultural fields is not always easy. There is a renewed interest in ensuring pollination services through practices that support pollinators. Pollinators like honey bees, birds, bats and insects, plays crucial role in flowering plant for the production of fruits and vegetables. Without the help of pollinators those plants cannot reproduce and they can pollinate two-thirds of our food crops (Anonymous, 2009h cited by Anonymous, 2009g). The countries like Canada, Germany, Italy, Japan, France, Great Britain, Australia, New Zealand, China, Argentina, Mexico, India, Netherlands, Korea, developed the beekeeping industry, and increased crop production tremendously. In Israel the beekeepers are able to commercialize the culture of bumblebees for crop pollination, particularly for greenhouses and they export these bees throughout the world. The fact is there, is no country on this planet that has productive crops without a developed beekeeping industry(Baconawa, 2009). The crops and pollinator species are belongs to each other and the world’s food supply depends on the health of these pollinators. There are more than 100,000 different animal species or perhaps as many as 200,000 species to playing a role in pollinating the 250,000 types of flowering plants on this planet. The annual monetary value of pollination services in the global agriculture industry could be as high as $200 billion (Anonymous, 2009i cited by Anonymous, 2009g). Honey bees pollinate approximately $10 billion worth of crops in the United States annually (Anonymous, 2009h, Watanabe, M.E. 1994 cited by Anonymous, 2009g); however, of the crops that make up most of the world's food supply, only 15% are pollinated by domesticated bees, while at least 80% are pollinated by wild bees and other wildlife (Anonymous, 2009h, Prescott and Prescott-Allen. 1990; Ingram, et al. 1996b cited by Anonymous, 2009g). Australian farmers rely heavily on the introduced Western honey bee to pollinate their crops. Trigona species (Stingless bees) and crop production The plants and the bees are made for each other because of their development are closely related. If the bees are allowed to cross-pollination, the crops are able to increase up to 40% in production. The bees are absolute pollinators compared to bats, birds; among others due to their maximum numbers for example Trigona colony has a maximum of 100,000 workers while Apis mellifera could have 60,000 maximum numbers of workers (Baconawa, 2009). The stingless bees have been shown to be valuable pollinators of crops such as macadamias and mangoes (Anonymous, 2009d). The pollination effectiveness of two species of stingless bees (Tetragonisca angustula Illiger and Nannotrigona testaceicornis Cresson) and Africanised honey bees (Apis mellifera L.) was determined for the ornamental plant Salvia farinacea Benth. All three species of bees highly increased the seed production in comparison with pollinator-deprived plants (Sanchez et al. 2001). They may also benefit strawberries, watermelons, citrus, avocados, lychees and many others. In the Philippine the Trigona spp.are better pollinators than the Apis species because they are very efficient and effective under this conditions. Pollinator bees, particularly Trigona, have great markets in Japan and Korea. These countries are now importing Trigona species from Australia. The Japanese are now appreciating the value of Trigona over the Apis species because of they are worry of the stinging behaviours of the Apis spp. Trigona have very low tendency to swarm because of more than two queens per colony, also rare to absconding and longer working life up to 60 days/worker. Trigona bees’ workers foraging habit up to 500 meter radius it means more intensive pollination of crops near hive. Also they have very short tongue so they can gather more nectar and pollen compare of other species. Trigona species are highly tolerance to the pest and diseases because of their smaller size and extensive use of propolis which serves as germicidal and pest repellent also they can highly tolerance to heat because of their

propolis canopy (Baconawa, 2009). Due to above mentioned characters, Trigona species are the best pollinator for pollination in horticultural crops.

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