Amegilla+zonata

// Amegilla zonata // (Linnaeus, 1758)  Blue-banded bee

__**Introduction**toc __
When one thinks of bees, the image that comes to mind is that of the typical honey bee, with its trademark black and yellow abdomen. However, there is a large variety of bees that do not follow this form, of which many can be found in Singapore. One such example is the Blue-banded bee (//Amegilla zonata//) which, as its name suggests, has blue bands on its abdomen instead of yellow.  “Blue-banded bee” can actually refer to many different species within the genus //Amegilla// which share this characteristic blue banding. //Amegilla// spp. are found strictly in the Old World 1], ranging throughout Australia, Asia, Europe and Africa, with currently 252 species described 2]. They are largely similar to and can be confused with bees of genus //Anthophora//, but //Amegilla// is easily differentiated by its lack of arolia, which are padlike projections between the tarsal claws 3].  Due to the relatively large distribution and snumber of species in //Amegilla//, it is further divided into 11 subgenera. //Amegilla// //zonata// is placed in subgenus //Zonamegilla//, and is the type species of this subgenus [1 ]. Through this webpage, I hope to be able to showcase a remarkable and unique bee species in Singapore, and show that not all bees are like the more commonly known honey bee.

**__Name __**
Binomial name: //Amegilla zonata// Vernacular name: Blue-banded bee Synonyms: //Apis zonata // Linnaeus, 1758 //Anthophora zonata // (Bingham, 1897; Friese, 1918) //Anthophora zonata // var. //puttalama// (Strand, 1913) //Amegilla (Zonamegilla) zonata //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> (Brooks, 1988)

**__<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Etymology __**
<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The species name //zonata// is derived from the Latin “zonatus”, which means either “belted” or “zoned”. This likely refers to the distinct metallic-blue bands on its abdomen. Its common name directly refers to this feature on the abdomen as well.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Diagnosis
<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The following characteristics can be used to differentiate //Amegilla zonata// from other bees, whether in //Amegilla// or other genera [ <span style="font-family: 'Times New Roman',serif; font-size: 16px;">1 <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">]4]. For reference, a diagram showcasing the morphology of a typical bee can be found [|here].
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">11-13mm in length.
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Seven exposed metasomal terga,
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Apical margins of abdominal segments 1-4 covered in metallic blue hair (see Fig 2)
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Yellow markings along labrum, clypeus, base of mandibles, and medial line across head (see Fig 3)
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Brown, short hairs on thoracic segments <span style="font-family: 'Times New Roman',serif; font-size: 16px;">(see Fig 3)
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Apex of final abdominal segment bilobed
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Apex of (male) gonostyle quadrate, bent inwards <span style="font-family: 'Times New Roman',serif; font-size: 16px;">(see Fig 4)
 * [[image:Amegilla_zonata_front_TelokBlangahHillPark_DavidTan_07Nov13.jpg width="496" height="340"]] || [[image:taxo4254/Amegilla_zonata_gonad_GBTB_JWTTan_08Mar14.jpg width="469" height="342"]] ||
 * Fig 3. Yellow markings across the head, brown short hairs on thorax (Photo: Chua Si Hao) || Fig 4. Quadrate gonostyle on the male (Photo: Chua Si Hao) ||  ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Comparison with //A. korotonensis//
//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla zonata //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">shares many resemblances to other //Amegilla// spp., particularly those in subgenus //Zonamegilla//. In Singapore, we are able to find one other species of //Amegilla//, namely //Amegilla korotonensis//. Fortunately, these two species are relatively easy to differentiate through the following characteristics:
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Characteristic ** || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">A. zonata // || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">A. korotnensis // ||
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">**Size** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Relatively larger || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Relatively smaller ||
 * **<span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Abdomen ** || <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Relatively larger || <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Relatively smaller, rounder ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Blue bands ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Darker, more metallic shade of blue || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Lighter shade of blue ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Hind legs ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">White hairs stretch from hind femur to tibia || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">White hairs only found on tibia ||
 * [[image:Amegilla_zonata_DairyFarm_ZestinSoh_05Jun12+korotonensis_HortPark_ZestinSoh_18Jun12_TOP.jpg width="800" height="533"]] ||
 * Fig 5. //A. korotonensis// (left) and //A. zonata// (right). We can see the difference in size and abdomen banding in this comparison. (Photo: Chua Si Hao) ||
 * [[image:taxo4254/Amegilla_zonata_DairyFarm_ZestinSoh_05Jun12+korotonensis_HortPark_ZestinSoh_18Jun12_SIDE.jpg width="800" height="533"]] ||
 * Fig 6. //A. zonata// (left) and //A. korotonensis// (right). The difference in white hairs on their hind limbs is clear here. (Photo: Chua Si Hao) ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Description
<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Original description by Carl Linnaeus (1758) 5]: “A. subpubescens fusca, abdomine cingulis quatuor caeruleis. Habitat in Indiis.”, which translates to “brown, short pubescence; four blue abdominal belts. Habitat in Indies”.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Feeding
(Photo: Khew S K, used with permission) || Fig 8. //Amegilla zonata// showcasing its long proboscis (Photo: Zestin Soh, used with permission) ||
 * [[image:Flower-and-bee.jpg width="458" height="312"]] || [[image:13242527803_736a03f2fb_o.jpg width="460" height="313"]] ||
 * Fig 7. //Amegilla zonata// performing buzz pollination on a //Melostama malabathricum// flower

//<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Amegilla zonata //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> employs buzz pollination in order to obtain pollen from flowers, a method not employed by honey bees. This involves grabbing onto the anthers of the flowers and vibrating the wing muscles at the right frequency to dislodge the pollen 6]7]. We can see an example of buzz pollination in Video 1. The pollen is then gathered on the hairs of the hind legs of the bee8]. //A. zonata// is also able to use its relatively long proboscis to feed on nectar at the base of the flower 9], which can be seen in Video 2.

//<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Amegilla zonata //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> is a trap-lining bee [ <span style="font-family: 'Times New Roman',serif; font-size: 16px;">9 <span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">], which means that it regularly visits the same flowers for food. In Singapore, //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">A. ////<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">zonata //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> can be observed visiting flowers such as //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Melastoma ////<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">malabathricum //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> (Fig. 9), //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Asystasia gangetica //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> 10], and //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Galphimia ////<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">glauca //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> (Fig. 10). They forage diurnally, but can be found more frequently during the late morning hours [ <span style="font-family: 'Times New Roman',serif; font-size: 16px;">6 <span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">]. (Photo: Zestin Soh, used with permission) || Fig 10. //Amegilla zonata// with //Galphimia glauca// (Photo: [|J. S. Ascher]) ||
 * media type="youtube" key="xD-wrZ3ZUWo" width="435" height="250" || media type="youtube" key="kws65tIt5Ps" width="420" height="315" ||
 * Video 1. //Amegilla// sp. performing buzz pollination || Video 2. //Amegilla// sp. using its proboscis to feed on nectar ||
 * [[image:taxo4254/13242376785_95bd2e1491_o.jpg width="435" height="297"]] || [[image:taxo4254/Amegilla_zonata,_anthophorine_bee,I_JSA4072.jpg width="319" height="307"]] ||
 * Fig 9. //Amegilla zonata// with //Melostama malabathricum//

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Ecosystem service **
//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> //zonata// serves an important role in most ecosystems by pollinating various flowers. It has been found to be especially important in tropical rainforest understories, which lack the large quantities of eusocial bees normally found in forest canopies [ 9 ]. Bees like //A. zonata// are particularly important as their target flowers may not be able to be pollinated without this buzz pollination. Many of these flowers even form specific relationships with their respective pollinators, further highlighting the importance of // A. //// zonata // in such forest settings.

//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">spp. have also been found to be effective at pollinating crop plants, such as tomatoes, under greenhouse settings. Such an ecosystem service would allow us to cultivate certain crops in Singapore and slightly alleviate our reliance on importing food. This also removes the need to import potentially invasive bees, such as honey bees or bumblebees, as //A.// //zonata// can be readily found in Singapore 6 ].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Nesting
<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">While the honey bees typically construct nests with hexagonal honeycomb cells, Blue-banded bees do things differently. Like most Anthophorine bees, //Amegilla// //zonata// builds its nest by burrowing into the soil 3]. While they are solitary bees, they often nest together in nearby burrows 11]. Various bees of //Amegilla// spp. may even be seen nesting together 1]. It has been suggested that pheromones released from the Dufour’s gland into the excavated soil may attract other females to nest 12]13]. Aggregated nesting appears to lend an advantage against destruction, predation, or parasitism 14]. Individuals have even been found returning to successful nesting sites of previous generations 15], highlighting the relative advantage such aggregations provide.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The burrows branch into individual cells, which are lined by secretions from the [|Dufour’s gland] that later solidify into a waxy layer 16]. The female then lays an individual egg into each cell, depositing it onto a provision of nectar and pollen. The female then closes up the cell with more of the waxy lining. The nectar, pollen, and even the waxy lining act as food for the larvae when it emerges. We can contrast this with the honey bee, which uses the nectar it collects to create honey, which is then fed to its larvae <span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">3]. <span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">One might argue that the pollen and nectar mixture that //A. zonata// produces reflects a rudimentary form of honey production in Anthophorine bees. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Males, on the other hand, do not nest in burrows, nor do they assist in the excavation. Instead, males rest on plant stems by grabbing on with their mandibles 17]. We can see this behavior in Video 3.
 * media type="youtube" key="2ia-SiSuKLI" width="420" height="315" ||
 * **Video 3.** Male //Amegilla// sp//.// roosting on grass stems ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Associated parasites
//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> bees are known to be targeted by kleptoparasitic //Thyreus// spp., commonly known as Cuckoo bees 18]3]. In Singapore, //A.// //zonata// faces parasitism from //Thyreus// //himalayensis// and //ceylonicus//19]20]. //Thyreus// bees break into the cells of the host nest, oviposit in it, and seal it back up with earth. The larvae then hatch and take advantage of the food deposited by the female //A.// //zonata// 3 ]. // Thyreus // bees also possess blue markings and bandings on their abdomen, which may serve as a form of camouflage against // Amegilla //. This has led some to propose that // Thyreus // may have evolved from a common ancestor as // Amegilla //, hence their similar colouration and associations, although this hypothesis lacks strong evidence thus far 3 ].

(Photo © www.NatureLoveYou.sg, used with permission) || Fig 12. //Thyreus ceylonicus//, which can be differentiated from //T. himalayensis// by a break in the horizontal band of blue hair across the abdomen (Photo © www.NatureLoveYou.sg, used with permission) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Besides //Thyreus// spp., //Amegilla// spp. have been observed to be parasitized by insects from other Orders, such as Diptera (//Miltogramma// sp.) and Coleoptera (//Zonitis// sp., //Trogoderma// //apicenne//, //Lusioderma// //serricorne//). Some cells have even been found with fungal colonies that feed on the deposited food and the larvae itself 18].
 * [[image:taxo4254/DSC00993 (13).JPG]] || [[image:DSC03406 (13).JPG]] ||
 * Fig 11. //Thyreus himalayensis//

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Global distribution
//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> //zonata// have been described to be found throughout India, Myanmar, Sri Lanka, extending through the Malayan peninsula to Australia 4]. There have also been records as far north China and Taiwan, however these may be misidentifications as many //Zonamegilla// species share strong resemblance 21].
 * [[image:Map_of_amegilla_zonata.jpg width="800" height="400"]] ||
 * Fig 13. Map showing global distribution of //A. zonata// 22] ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Local distribution
//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Amegilla //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> //zonata// can be found at the locations in the below map. Data is obtained from National University of Science Bee Lab (collected by J. S. Ascher, S. H. Chua, S. X. Chui, K. Y. Ng, Z. W. W. Soh, E. J. Y. Soh, J. W. T. Tan, and K. S. Yow). <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">As we can see, //A. zonata// can be found at many parks and secondary forests ranging throughout Singapore, so keep a look out for them the next time you are in a park or our nature reserves.

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__<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Taxonomic classification __

<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Kingdom Animalia <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Phylum Arthropoda <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Subphylum Hexapoda <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Class Insecta <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Order Hymenoptera <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Family Apidae <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Tribe Anthophorini <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Genus //Amegilla// (Friese, 1897) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Subgenus //Zonamegilla// (Popov, 1950) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Species //Amegilla zonata// (Linnaeus, 1758)

__<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Phylogeny __
<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The phylogenetic relationship of various genera and subgenera under the Tribe Anthophorini is displayed below 23]. It showcases the close relationship between //Amegilla// and //Anthophora//, explaining their similar general morphology. It also shows a close but undefined relationship between subgenera //Zebramegilla, Glossamegilla//, and //Zonamegilla// (including //A. zonata//).



__<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">References __
>
 * 1) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Brooks, R. W. (1988). Systematics and phylogeny of the anthophorine bees (Hymenoptera: Anthophoridae; Anthophorini). //The University of Kansas science bulletin (USA)//.
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 * 12) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Cardale, J. C. (1968). Nest and nesting behaviour of Amegilla (Amegilla) Pulchra (Smith)(Hymenoptera: Apoidea: Anthophorinae). //Australian Journal of Zoology//, //16//(4), 689-707.
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 * 16) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Batra, S. W. T. (1972). Some properties of the nest-building secretions of Nomia, Anthophora, Hylaeus and other bees. //Journal of the Kansas Entomological Society//, 208-218.
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 * 18) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Cardale, J. (1968). Parasites and other organisms associated with nests of Amegilla Friese (Hymenoptera: Anthophorinae). //Australian Journal of Entomology//, //7//(1), 29-34.
 * 19) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Ascher, J. (n.d.). Thyreus ceylonicus (Friese, 1905). Retrieved November 4, 2014, from http://www.discoverlife.org/mp/20q?search= Thyreus+ceylonicus
 * 20) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Ascher, J. (n.d.). Thyreus himalayensis (Radoszkowski, 1893). Retrieved November 4, 2014, from http://www.discoverlife.org/mp/20q?search= Thyreus+himalayensis
 * 21) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Engel, M. S. (2007). A new Amegilla of the zonata group from Malaysia and Thailand (Hymenoptera: Apidae). //Transactions of the Kansas Academy of Science//, //110//(1), 16-22.
 * 22) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Ascher, J. (n.d.). Amegilla zonata (Linnaeus, 1758). Retrieved November 4, 2014, from <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[]
 * 23) <span style="color: #222222; font-family: 'Times New Roman',serif; font-size: 12pt;">Dubitzky, A. (2007). Phylogeny of the World Anthophorini (Hymenoptera: Apoidea: Apidae). Systematic Entomology, 32(4), 585-600.