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Int.J.Curr.Microbiol.App.Sci (2019) 8(7): 580-587

International Journal of Current Microbiology and Applied Sciences ISSN: 2319-7706 Volume 8 Number 07 (2019) Journal homepage: http://www.ijcmas.com

Review Article https://doi.org/10.20546/ijcmas.2019.807.072

A Review on Diversity, Bio-Ecology, Floral Resources and Behavior of Blue Banded

J. Sandeep Kumar1*, B, Rex2, S. Irulandi3 and S. Prabhu3

1Department of Entomology, Agriculture College and Research Institute, Madurai-625104, India 2Department of Plant Pathology, Agriculture College and Research Institute, Madurai-625104, India 3Department of Plant Protection, Horticulture College and Research Institute, Periyakulam-625604, India

*Corresponding author

ABSTRACT

K e yw or ds Blue banded bees are solitary bees which are characterized by their Blue banded bees, glittering blue color bands on their abdomen. They construct their nests and , Diversity, brood cells for their young ones in soils. The developmental biology of blue Behavior, India banded bees is maximum sixty days and varied according to climatic Article Info conditions. They are polylectic and wild pollinators playing major role in

crop pollination. Research on blue banded bees is very limited and less Accepted: 07 June 2019 studies. In this review we will focus on diversity, bio-ecology and behavior Available Online: of blue banded bees. 10 July 2019 Diversity of blue banded bees (Brooks, 1988; Michener, 2000). The two important genera of the tribe are Blue banded bees are Anthophorine bees. An and Amegilla which are easily Anthophorine was first described by distinguished based on the presence or Linneaus in 1758 as Apis retusa. Latreille absence of arolium. The male genetalia of described first Anthophorine in 1802 different Anthophorine bees vary distinctly and in 1803 he proposed the name (Brooks, 1988). They frequently bear metallic Anthophora. Fabricius named a genus as blue or green pubescent bands on metasomal Megilla in 1805. The name Megilla is now terga. They fly in summer. This genus regarded as a junior synonym of Anthophora includes 12 sub genera and 253 species by Michener, (1974). Anthophorine bees are (Brooks, 1988). The genus Amegilla is a now grouped under the family , sub diverse group of approximately 255 family and tribe Anthophorini Anthophorine bee species distributed in

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Int.J.Curr.Microbiol.App.Sci (2019) 8(7): 580-587 southern Europe and Mediterranean basin Amegilla in his revision of Egyptian southward throughout Africa and Madagascar Anthophora species Engel (2007) described east into Arabia and in Asia as far as northeast new species of Amegilla from northeastern China, India, Sri Lanka, Indonesia, New Egypt viz., Amegilla argophenax and Guinea as well as Australia (Engel, 2007). In differentiated same from the commonly the oriental region there is a large diversity of occurring species. Blue banded bees shows blue banded bee species that are frequently sexual dimorphism, opposite sexes can be confused with each other, typically being differentiated based on their body size where identified as the relatively widespread male bees are smaller than female bees, head . Closer examination of these appendages (Clypeal markings are absent in bees reveals that there are multiple distinct male bees, mandibles are short in male bees, species among Southeast Asian “zonata”. antenna is short in female bees than males, Engel (2007) described a new species of number of flagellomeres are eleven in male Amegilla from zonata group and named it as and ten in female, compound eyes are small in Amegilla anekawarna Engel from Malaysia. male bees and sting is absent in male bees) The new species is similar to A. zonata but and blue color metallic bands on metasoma. can be distinguished by the presence of a Gonostylus is lessen to a blister in male bee distinct medioapical blue band on fifth (Sandeep and Muthuraman, 2018). metasomal tergum and by the yellow ventral surface of scape in the female and presence of Nesting biology two dark clypeal markings in male. Rayment (1951) revised 15 Australian species of blue Several workers studied the soil nesting banded bees and grouped them under the behavior of Anthophorine bees (Thorp, 1969; genus Asaropoda. A. dawsoni is Australia‟s Torchio 1974; Brooks, 1983; Norden, 1984; largest Anthophorine bee. In these species Houston, 1991; sandeep and Muthuraman, two types of males i.e. large and small males 2018). Blue banded bees are fossorial bees are present. Males exhibit a bimodal size and build their nests underground (Cane, frequency distribution. Amegilla is the only 1981). Their nests are burrows in soil, either genus occuring in Australia where it is in banks or in flat ground (Michener, 1960). represented by three subgenera Asaropoda, The female bee also pays attention for the Notomegilla and Zonamegilla (Brooks, 1988). surface nature of nesting site and edaphic Lieftinck (1956) revised the oriental attributes of the soil (Cane, 1991). These bees Anthophorine bees of the genus Amegilla and build nests in a variety of soil types like black described sixteen species under these genus soil, clay soil and sandy soil but they always occuring both in Malaysia and Southeast prefer a site with some shelter (Rayment, Asia. Lieftinck (1975) redefined four species 1951). Anthophora edwardsii preferred of Amegilla complex viz., A.florea, A.confusa, crumbly clay-silt in the open areas to loose A. parhypate and A.calceifera in Korea. sandy silt in the vegetated areas (Thorp, Brooks (1988) proposed a subgeneric name 1969). Many soil-dwelling solitary bees make Glossamegilla for Amegilla sp. with long their nests in groups or aggregations where glossa collected from India to the Indo- females nest gregariously and a clumped nest Malayan regions and included 29 species in distribution is found (Batra, 1978).The female the group. Attasopa and Warrit (2012) bee prefers to build the nest at the nesting redescribed the subgeneric position and sites from where it emerged as an adult. This distribution of oriental burrowing bee, phenomenon is termed as philopatry. The Amegilla fimbriata. Michener (1956) included philopatry may be a probable reason for nest

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Int.J.Curr.Microbiol.App.Sci (2019) 8(7): 580-587 aggregations (Norden, 1984). In initiating a cells. Brood cells are cylindrical and urn new nest, a female selects a site and begins to shaped. Waxy coatings were found in inner excavate a burrow by biting at the soil with cell wall of brood cells which would be used her mandibles. The soil is periodically for water proof and larval feed, larval faeces moistened and softened with liquid from the and cast skin was found inside the brood cell. mouth. Females interrupt their burrowing Nests are often built together in one place from time to time and depart on flights away forming nest congregation and the maximum from the nesting area presumably to recharge number of nests was 78 per m-2 (Sandeep and their crop with liquid (Houston, 1991). Muthuraman, 2018). Ground nesting bees sometimes conceal their nest entrances in loose sand under clods or in Developmental biology dense vegetation. However, most of them leave some kind of mound or other tell-tale Blue banded bees are fossorial bees. The life mark at the entrance. The nests are shallow stages of the bees develop inside sealed seated inside the soil and usually less than 10 earthen brood cells. More often invasive cm deep. Excavated soil on horizontal methods are used for studying the life stages surfaces is often heaped in a small cone or of these bees. Of late X-ray computerized delta of irregularly sized pellets, near the nest tomography (CT) scanning is used to study entrance (Cane, 2008). Chimney bee the developmental biology and natural A.abrupta digs a tunnel for its nest and uses enemies of bees (Greco et al., 2006). As in all the excavated soil to build a chimney or turret Hymenopteran the sex is determined near nest entrance. Each turret corresponds to by haplodiploidy mechanism. The male bees a single nest (Norden, 1984). Turrerts give develop from haploid eggs and female protection to nest from rain, nearby develop from diploid eggs. The bee larvae are excavations, windblown debris or parasites. reared through mass provisioning. The larval They also function as a land mark for nest food may be a thick liquid or semi-solid in recognition and thermoregulation of nest consistency. Immature life stages of Amegilla, (Brooks, 1983; Norden, 1984). The burrow Asaropoda and Anthophora are similar in descends down as a shaft which ends in urn- morphology (Cardale, 1967). In A.dawsoni shaped terminal cell or a cluster of cells or in the female is capable of producing two kinds a linear series. The inside of the cell is of male‟s viz., namely major males and minor extremely smooth and polished and then males by constructing larger and smaller covered with a heavy layer of white, water brood cells and provisioning the cells with proof material may be flaked off from the cell different amount of food (Tomkins et al., wall made of Dufour‟s gland lipids (Thorp, 2001). Females of Dawson‟s burrowing bees 1969; Cane, 1991). The various substrates have a well-defined brood cell cycle suitable for nesting were studied under pot involving cell construction, waxing, culture experiment. Red soil and potting provisioning, egg laying and cell capping mixture were recorded for best substrates for (Alcock, 1998). Eggs are singly laid by the nesting. Mostly nests were found close to female bee on the larval provision (Cane, edge of pot. The female bee digs out her own 1984). Eggs hatched in about five days and nest burrow of depth 4 - 30 cm using her the larvae developed through four instars by strong mandibles and excavated soil by her eating the larval provision. The last instar legs forming a tumulus and some bees formed larva after consuming larval provisions turrets (1.2 to 4.8cm). The nest shaft extends consumed the cell linings and defaecated from entrance to a cluster of earthen brood within the cell which formed the basal lining

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Int.J.Curr.Microbiol.App.Sci (2019) 8(7): 580-587 of the cell and turned into prepupa without Floral resources moulting. They do not spin cocoons. Bees over wintered as prepupae under temperate Studies on foraging resources of blue banded conditions and completed one generation in a bees are scanty. The bee foraged on both wild year. In tropical conditions the developmental and cultivated flora and hence they arrest was not found. In A. abrupta egg period constituted an important component of the lasted for 5 days, larval period for 21 days pollinator guild of an area. The species of the and pupal period for 18 days (Norden, 1984). plant family‟s Verbanaceae, Lamiaceae, Cardale (1967) reported the natural enemies Acanthaceae were more preferred by these associated with Amegilla pulchra. He found bees for the collection of floral rewards. out a cleptoparasitic bee ( These bees were able to compete with other caeruleopunctatus), a Sarcophagid fly anthophilous insects in sharing floral (Miltogramma sp), a Meloid beetle (Zonitis resources. Flowers with white, purple, blue sp), a mite (Pymotes ventricosus) and a and yellow were more attractive for this bees. fungus (Aspergillus sp). Norden and Intra floral behavior of avoiding stigma Scarbrough (1982) studied about three contact while foraging on Moringa oleifera important parasites affecting Anthophora and Bauhinia purpurea reduced the chances abrupta. They recorded a Conopid fly of pollination. Amegilla was found to be a (Physocephala marginata) as an adult rapid flier and even few individuals covered a parasite. Green metallic cuckoo bee (Chrysis large number of flowers in their foraging trip sp) was parasitic on immature stages. A (Subba reddi et al., 1999). The foraging Meloid beetle (Hornia minutipennis) was behavior of a blue banded bee Amegiila found to feed on bee eggs and larval chlorocyanea was studied in Australian provisions. Houston (1991) found out a tomato greenhouses by (Hogendoorn et al., Sarcophagid fly (Tayloromyia iota) affecting 2007). Their study revealed the foraging the adult female bee, A.dawsoni. The potentiality and the practical utility of blue endoparasitic maggots killed the parasitized banded bees for pollinating tomato grown in bee. Eggs are laid individually on bee bread greenhouses instead of using the exotic (0.07 g) in the brood cell was mass bumble bees. provisioned by the bees. The first and second instar larva was straight and slightly They were active foragers and performed on pigmented mandibles respectively. Third an average nine pollen flights per day when instar larva was „C‟ shaped along with small foraging in a tomato greenhouse. The bees amount of unutilized brood food. The fourth collected pollen from the anthers by shaking instar larva almost occupies the brood cell the flowers in flight (Hogendoorn et al., without any food materials. Larva consumes 2006). The flight activity of both male and wax linings and shrinks slightly before female bees started early morning around turning into a prepupa. Pupal eye coloration 6:00am ends during sunset at 6:30pm and changes from yellow to orange and then to collect floral nectar or pollen. These bees brown and finally black as the pupa matures. preferred various plants from Fabaceae, The adult bee emerges fifteen days after Solanaceae and Cucurbitaceae. These bees pupation. It was found that the development collected pollen or nectar from12 weeds and of bees were pretentious by heavy rains and 21 crop plants and they are attracted mainly to brood mites (Rhizoglyphus sp. and yellow coloured flowers followed by white Histiostoma sp.) (Sandeep and Muthuraman, and purple colors (Sandeep and Muthuraman, 2018). 2018).

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Behaviour not mount them. It is not known if contacts between males were a result of mistaken Male bees were found in both nesting areas identity or aggression (Norden, 1984). These and at forage sites. At nesting areas, they fly brief encounters resulted in both males slowly over the ground (within 15 cm of the departing. Similar encounters between male surface) following meandering paths. They Anthophora occidentalis were noted by Batra paid particular attention to holes without (1978). Sleeping aggregations have been turrets which they frequently inspect and observed in various solitary bees by Michener sometimes entered briefly. They pay no (1974). Such aggregations consist of usually attention to turreted burrows or females conspecific males rarely with few females entering or leaving them. The male bees wait gathering on plant stem. In most of the at the nest entrance for the virgin female bee Hymenopteran species males have been to come out. They can detect the presence of observed spending the night on flowers or female bee even while she is inside the tunnel. leaves but most of the females spend the night The male bees fight with each other using in burrows or nest structures. However, males their mandibles and winning bee alone can of several solitary bee and wasp species have mate with female bee (Houston, 1991). observed aggregating in a specific location Mating occurs at nesting areas and mainly at overnight. Female biased aggregations of the forage sites. The female bee normally Amegilla florea were reported by Yokoi and mates only once and mating lasts about seven Watanabe (2014) in Japan. The sleeping minutes in (Simmons et al., aggregation observed was unusually female 2000). The female, mounted by the successful biased and occurred on dead branches. The male, ran over the ground for some meters sex ratio and number of sleeping individuals until the pair was hidden beneath some small in the sex mixed aggregation changed daily. plants and there coupling lasted for about two Female arrived later but left earlier than male minutes (Houston, 1991). Males conducted to avoid long stay with male. The sleeping apparent searching maneuvers by darting sites were shared by both the sexes for want from flower to flower and hovering above of enough places to sleep. Aggregation each blossom for one to two seconds. They behaviours have been reported in males since occasionally stopped to drink nectar, but then the 1800s. Target locations and substrates for resumed their flights. Mating was initiated sleeping aggregations include branches, plant after a male hovered for approximately one stems, leaves and flowers. Individuals often second above a feeding female. He flew return to the same site in the late afternoon or rapidly to the female, butted her abdomen evening, reforming the aggregation in the with his head, and then grasped her thorax same location for weeks, months (Linsley, with his fore and mid legs. While in a dorsal 1962; Alvis-dos-santos et al., 2009). position, his genitalia were inserted and his Gregarious sleeping was observed in Amegilla abdomen began a pumping motion. Male garrula in Italy by Aldini (1994). The mouthparts were sometimes extended and aggregation had both male and female. stroked the female's head. Copulation lasted However no mating occurred at the between 15 and 45 seconds with females aggregation site. Female bee guards the nest remaining passive throughout in A.abrupta by remaining within the nest shaft. They (Norden, 1984). Previously mated females hover around the intruder at nest congregation were occasionally mounted by other males, site but they do not sting. They sting only but always avoided copulation. Males also when provoked or roughly handled at nesting, commonly butted other feeding males but did foraging and sleeping sites without losing

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Int.J.Curr.Microbiol.App.Sci (2019) 8(7): 580-587 their sting. Most of the sleeping clusters are : Apidae). Gentetics intraspecific and male biased and rarely sex Molecular Research. 8, 515-524. mixed. Males take rest during dusk either Attasopa, K., and Warrit N. Redescription, singly or in a cluster on the terminal end of subgeneric position and distribution of the dried weeds (Achyranthes aspera, Chloris the Oriental burrowing bee, Amegilla barbata and Vicoa sp) by grasping the stem fimbriata (Hymenoptera: Apidae: firmly with their mandibles and fall asleep. Anthophorini). Pan-pacific Entomology. Interspecific sleeping clusters are also found 88 (3): 281-291. occasionally which involved cuckoo bees Batra, SWT, 1978. Aggression, Territoriality, (Thyreus sp.) and leaf cutter bees (Megachile Mating and Nest Aggregation of some sp.). When both the sexes are brought Solitary Bees (Hymenoptera: together and confined in enclosures forced Halictidae, Megachilidae, Colletidae, mating can occur (Sandeep and Muthuraman, Anthophoridae). Journal of Kansas 2018). Entomologocal Society. 3 (51): 547- 559. India has a very large bee fauna, but often Bell MC, Spooner-Hart RN and Haigh AM. threatened by agricultural practices and 2006. Pollination of greenhouse natural calamities. There few basic tomatoes by the Australian blue banded information about biology and behavior of bee Amegilla (Zonamegilla) holmesi blue banded bees in India. Studies on Rayment (Hymenoptera: Apidae). distribution and taxa of blue banded bees are Journal of Economic Entomology. 99, very limited. Hope this review will encourage 437–442. more researchers to work on these bees. Brooks, RW, 1983. Systematics and Future work on developmental and pollination bionomics of Anthophora: The studies will help farmers to employ a cost Bomboides group and species group of effective pollinators in their ecosystem. the new world. University of California Publications in Entomology.98, 1-84. References Brooks, RW, 1988. Systematics and phylogeny of the anthophorine bees Alcock, J., 1998. Sleeping aggregations of the (Hymenoptera: Anthophoridae; bee Idiomelissodes duplocincta Anthophorini).University of Kansas (Cockerell) (Hymenoptera: Science Bulletin. 53(9): 436-575. Anthophorini) and their possible Buchman, SL., 1992. Buzzing is necessary for function. Journal of Kansas tomato flower pollination. Entomological Society. 71, 77-84. Bumblebeequest. 2, 1–3. Aldini, RN., 1994. Preliminary observations Butler, CG, 1965. Sex attraction in Andrena on gregarious sleeping in Amegilla flavipes Panzer (Hymenoptera, Apidae) garrula (Hymenoptera: Apoidea: with some observations on nest-site Anthophoridae). Ethology Ecology and restriction. Proceedings of Royal Evolution. 3,131-135 Entomological Society. 40, 77-80. Alvis-dos-santos I, Gaglianone MC, Naxara Cane, JH, 1981. Dufour's gland secretion in SRC and Engel MS. 2009. Male the cell linings of bees (Hymenoptera: sleeping aggregation of solitary oil Apoidea). Journal of chemical collecting bees in Brazil (: Ecology.7, 403-410. Tapinotaspidini and Tetrapediini: Cane, JH, 1991. Soils of ground-nesting bees (Hymenoptera: Apoidea): texture,

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How to cite this article:

Sandeep Kumar, J., B, Rex, S. Irulandi and Prabhu, S. 2019. A Review on Diversity, Bio- Ecology, Floral Resources and Behavior of Blue Banded Bees. Int.J.Curr.Microbiol.App.Sci. 8(07): 580-587. doi: https://doi.org/10.20546/ijcmas.2019.807.072

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