Tetragonula carbonaria
Tetragonula carbonaria | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Apidae |
Clade: | Corbiculata
|
Tribe: | Meliponini |
Genus: | Tetragonula |
Species: | T. carbonaria
|
Binomial name | |
Tetragonula carbonaria Smith, 1854
| |
Synonyms[1] | |
Trigona angophorae Cockerell, T.D.A. 1912 |
Tetragonula carbonaria (previously known as Trigona carbonaria
Tetragonula carbonaria forms honeycombs in their nests.[7] The bee produces an edible honey; the whole nest is sometimes eaten by Indigenous Australians.[8] The bees "mummify" invasive small hive beetles (Aethina tumida) that enter the nest by coating and immobilising the invaders in wax, resin, and mud or soil from the nest.[9]
Stingless bees
Twenty-one genera of stingless bees (family
Taxonomy and phylogeny
The eusocial stingless bees (Apidae, Apinae, Meliponini) comprise about 374 species.[11] Two genera occur in Australia, with Tetragonula being one of them.[11] The Tetragonula species of Australia were once in the larger genus Trigona, but were moved into a new genus in 2013.[12] Only minor structural differences are seen at the species level of the genus Tetragonula. T. carbonaria is nearly identical to T. hockingsi, besides a few differences in their nest architecture.[11]
Description
Bees
Compared to other stingless bees, T. carbonaria tends to be medium-sized.[6] However, their size can vary based on location. For instance, T. carbonaria from Queensland can be as small as T. mellipes, but in New South Wales, they can get as large as T. hockingsi.[6] They are all predominantly black-bodied, covered in microscopic hairs.[10] The adult workers and males are all black, with some brownish tint in certain areas such as the legs. The worker’s body length is 3.9–4.3 mm (0.15–0.17 in) and the wing length (including tegula) is 4.1–4.6 mm (0.16–0.18 in). The male drones have very similar bodies and wing lengths, but can be identified by different antennae.[6] T. carbonaria is a very strong flier, being able to fly up to 1 km (0.62 mi).[10] However, the species will not fly any further than they have to, so close resources are preferred over those farther away. T. carbonaria and its closely related species have high levels of morphological similarities. Also very low genetic variation exists within T. carbonaria.[13] T. carbonaria can be distinguished from other species of Australian Native Stingless Bees by their brood (in a distinctive spiral unique to the species), and by the entrance (these bees tend to daub resins around their entrances, where as other species, such as T. hockingsi generally keep their entrances clean.
Nest structure
The entrance and surrounding areas of their nest are coated with a smooth, thick layer of black, red, or yellow
Distribution and habitat
The nests are found in open forests and woodlands. They are usually built in tree cavities, and have small cryptic entrances, with no external entrance tube.[16] Four or five workers are usually visible at the entrance and are expected to be guards. They tend to choose larger trees and wider cavities to produce insulation valuable for their survival in the cool regions. Some features that would favour survival in a cooler climate are a high tree height and large feeding pots. The nesting sites of T. carbonaria are located near the top tree trunks that are 1.5 m (4 ft 11 in) in diameter, and are predominantly found in trees that are well insulated. Members of T. carbonaria also create the largest honey and pollen pots compared to the other species of the genus Tetragonula, which may help with efficient food storage.[6] In urban and suburban areas, T. carbonaria have been found to nest in Telstra pits as well as water meters when other habitat is not available.
Colony cycle
Activity
Colonies of T. carbonaria tend to be active all year round. The daily activity period, however, is longer in the southern hemisphere's warmer months, October to March.[3] The intensity of these daily flights is greatest in September, and least intense in May. A temperature threshold exists on all of this activity. Flight can only occur at temperatures greater than 18 °C (64 °F).[3] This year-long period of activity is beneficial for the pollination of crops flowering at any time of the year.
Initiation
Each brood cell is stocked almost to the brim with honey and pollen. An egg is laid in the cell by the queen and then the cell is closed. Complete larval and pupal development occurs in the closed cell .[10] Once the adult emerges, the cell is destroyed. Most stingless bee species are monogynous, meaning that when the colony divides, one of the daughter colonies will be queenless.[18] T. carbonaria colonies are frequently divided by beekeepers to increase the number of colonies. They build emergency queen cells by fusing two worker-sized cells that contain eggs or young larvae.[10]
The queens cannot live alone and they are not transferred to a new nest until it has been fully prepared by workers. The new queen is the bee that makes the flight to the new nest, with the old queen remaining in the parent nest.[10] When the old queen has died, mating swarms can occur at the established nest to replace the old queen with a young, unmated one.[18]
Growth
A study performed by Tim Heard in 1988 observed the propagation of hives in T. carbonaria. He successfully transferred colonies to boxes, and then once the available space was occupied, he would split the box by prying apart the two halves of the box. He recorded that colony weight increased much more in spring and summer compared to autumn and winter. After about 17 months, the final weight was established.[15] The rate at which colony weight increases is dependent on the availability of nectar and pollen, not age (however, it usually takes 12 to 18 months for a hive to be ready to be propagated). A heavy hive suggests filled storage pots and a large population of workers and brood, meaning the hive is ready to be split.[15]
Behaviour
Division of labour
Division of labour exists among the workers. The young bees perform tasks within the nests, such as brood care. As they mature, they become foragers and their tasks are performed outside of the nest.[19]
Foraging behaviour
T. carbonaria depends on
Reproduction
A study used
Kin selection
The workers tend to be the progeny of a singly mated queen. The colonies are predominantly
Worker-queen conflict
When workers do lay eggs, direct conflict tends to occur within the colony between the queen and the workers over the source of male eggs. Queen-worker conflict is found in cell provisioning and the
Interaction with other species
Defence
When the colonies are attacked, nest defence relies on the ability to recognise intruders. T. carbonaria sometimes displays a behaviour known as a "fighting swarm" when a non-nest mate is encountered.[16] Thousands of workers gather together and form a cloud. The signal to form this cloud is most likely mediated by alarm pheromones, which workers release from their mandibular glands.[16] As one entity, they drop to the ground and wrestle the intruders, which often leads to death of both parties. This behaviour is also a common defence mechanism against large predators such as humans.[16] T. carbonaria bees are highly sensitive to intruders, since they will even attack invaders that are carrying pollen or nectar. Even if an intruder found a way to make it past the swarm, it still would not make it through the congested entrance tunnel.[16]
Predators
One predator known to attack T. carbonaria is an Australian crab spider,
Parasites
The braconid subfamily Euphorinae has several genera, including Syntretus, known to be parasitoids of the adult stage of insects.[19] They are a highly diverse group and tend to be very successful parasitoids worldwide. A new species of Syntretus, S. trigonaphagus, has recently been discovered as parasitizing workers of T. carbonaria. Females of S. trigonaphagus are frequently found at the entrances of T. carbonaria hives near Queensland, Australia.[19] They approach workers that land nearby and oviposit on the host by curling their abdomens. The workers repeatedly brush their abdomens afterwards, suggesting that they were aware that an attack occurred. The overall effect of this parasitism is usually fatal. Older workers are more likely to be parasitized.[19] Because of this, as long as the number of parasites is minimal, the overall cost to the colony is not large, since these workers have already contributed substantially to the colony’s welfare.
Importance to humans
Beekeeping
Meliponiculture is the practice of stingless beekeeping,[24] where beekeepers maintain, reproduce, and use stingless bee colonies for their own profit. These colonies tend to be managed through artificial hives, so that the beekeepers have the ability to propagate the colonies and produce hive products such as honey and pollen. These products are then sold to various buyers from health-food stores to gift shops. The honey of T. carbonaria possesses a peculiar smell which makes it quite the appealing product. When the first work began in 1984 on this stingless bee, the industry was practically nonexistent.[24] Since then, the interest in stingless bees, more specifically T. carbonaria, has greatly increased. This has allowed for the establishment of conservation groups along the eastern regions of Australia. T. carbonaria is the most popular species that beekeepers tend, followed by the A. australis and then T. hockingsi. The main reason for most people to keep T. carbonaria is for enjoyment and conservation.[24] With this, the amount of honey produced is constantly increasing at a fast pace. T. carbonaria is the main species from which beekeepers harvest honey, of the stingless bees in Australia. Beekeepers report that one of the major limiting factors in propagating colonies is the availability of queens. More research is needed on queen rearing to fix this limiting factor.[24]
Agriculture (as pollinators)
Individual T. carbonaria bees demonstrate a consistency in floral choice. Individuals restrict their foraging activity to one kind of flower during a particular trip.[25] This consistency in a single pollen type enhances the pollinator efficacy by increasing the chances of pollen being transferred to stigmata of the same plant species. This increases their importance ecologically as crop pollinators. At the level of the colony, however, the species can use many different flowering species. So although the species is polylectic, individual bees remain consistent with their flower choice.[25]
Honey
Meliponines store their honey in pots, not in combs like the honey bees.[26] Compared to the honey of the Western honey bee A. mellifera, T. carbonaria honey had higher values in moisture, water activity, and electrical activity.[26] The two different honeys can also be distinguished by flavor and aroma. Also, the antioxidant activity of T. carbonaria honey has nutritional and pharmaceutical potential.[26]
References
- ^ a b Dollin, A.; Walker, K.; Heard, T. (2009). "Trigona carbonaria Sugarbag bee". PaDIL (Australian Biosecurity). Archived from the original on 1 April 2012. Retrieved 29 October 2011.
- ^ .
- ISBN 978-90-5410-486-5.
- S2CID 84566309.
- ^ doi:10.1071/it96020.
- ISBN 978-0-8018-6133-8.
- ISBN 978-0-415-92467-2.
- ^ "Australian bees 'mummify' their beetle enemy alive". BBC News. 17 December 2009. Retrieved 23 December 2009.
- ^ a b c d e f g h Heard, T; et al. (1996). "Stingless Bees". Nature Australia: 51–55.
- ^ S2CID 33790632.
- ISBN 978-1-4614-4960-7, retrieved 2022-06-28
- .
- ^ "Stingless Bees". ABeeC Hives - Australian Native Bee Hives. Retrieved 2022-07-26.
- ^ .
- ^ S2CID 44720135.
- ^ "OATH Beehives". ABeeC Hives - Australian Native Bee Hives. Retrieved 2022-07-26.
- ^ S2CID 83579207.
- ^ .
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- ^ S2CID 25563806.
- ^ S2CID 24236958.
- ^ S2CID 86553724.
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Further reading
- Dollin, Anne (December 2008). "The Mysterious Fighting Swarms" (PDF). Aussie Bee Online.
- Gloag, R.S.; Beekman, M.; Heard, T.A.; Oldroyd, B.P. (2008). "No worker reproduction in the Australian stingless bee Trigona carbonaria Smith (Hymenoptera, Apidae)" (PDF). Insectes Sociaux. 54 (4): 412–417. S2CID 25563806.
- "A New Name for our Trigona Stingless Bees: Tetragonula". Aussie Bee Online.
- Michener, C. M. (2013). "The Meliponini". In Patricia Vit; Silvia R. M. Pedro; David Roubik (eds.). Pot-Honey: A legacy of stingless bees. New York: Springer. pp. 4–8. ISBN 9781461449607.