Allodapini | |
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Allodapula sp. in South Africa | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Apidae |
Subfamily: | Xylocopinae |
Tribe: | Allodapini Cockerell, 1902 |
Genera | |
Allodape Contents |
The Allodapini is a tribe of bees in the subfamily Xylocopinae, family Apidae. They occur throughout sub-Saharan Africa, South East Asia, and Australasia. [1] There is also a rare genus, Exoneuridia, that occurs in isolated regions of Turkey, Iraq, Lebanon and Iran. [2]
Many of the species in the tribe form small social colonies where a group of females cooperatively care for the developing larvae. [3] The larvae are fed on pollen which, like most other bees, is carried on specialised hairs of the hind pair of legs, but the pollen is fed to the larvae in a progressive fashion and usually placed directly onto their bodies where they then consume it.
The larvae of allodapine bees are remarkable in their complex morphology, and in most species they possess appendages, tubercles and long setae. [4] The strange morphology of allodapine larvae is probably a result of living in open tunnels where they are in constant contact with other larvae and with adults. The appendages, tubercles and setae serve to hold and manipulate food, and may also help larvae move around the nest. These abilities are important because larvae compete with each other to gain food, a situation which is different from all other bees, where individual larvae are isolated in cells and do not have to compete with each other.
There are over 300 described species of allodapine bees, [1] but many more species are undescribed. They are unique among bees in progressively rearing their larvae in undivided tunnels, so that individual larvae are not physically isolated from each other and are in constant contact with adult females, who provide them with food, groom them, and remove their faeces. [1]
Allodapine bees vary greatly in their forms of sociality, from subsocial to highly eusocial. [5] There are no known species that are purely solitary. [6] They have been used widely to study social evolution, [7] sex allocation, [8] social parasitism, and historical biogeography. [9]
Many allodapine species exhibit very simple forms of social organization, without clear queen or worker castes. For this reason it was long thought that they had only recently evolved forms of social living. [10] However, molecular phylogenetic studies show that social living is ancestral for the tribe as a whole and has been in place for about 50 million years. [9] An ancient origin of sociality in this group helps explain very sophisticated forms of social communication in some species, such as pheromonal regulation of reproduction [11] and complex forms of kin recognition. [12] The origin of queen and workers castes in allodapine bees is relatively recent, much less than 40 million years ago, compared with the honeybees, bumble bees and stingless bees, where true queen and worker castes evolved about 100 million years ago. [13]
Most allodapine bee species have strongly female-biased sex ratios, and in many species less than 15% of brood are male. [8] This is very different from the vast majority of animal species where sex ratios are very close to 1:1 males:females. The preponderance of female-biased sex ratios in allodapine bees is thought to be due to the benefits of sisters cooperating with each other and involves a theory known as local resource enhancement. [14] For example, in Exoneura robusta , females provide the useful work in the colony and group living increases colony success, so the sex ratio is almost always female biased in this species. [15] [16]
Socially parasitic allodapine bees are species that have evolved to exploit the social systems of their hosts (which are other allodapine bees) so that the parasites enter the host colonies and lay their eggs there, and both the parasite adults as well as their larvae are fed by the host species. Molecular research has revealed nine origins of social parasitism in allodapine bees, [17] more than all other bees and wasp groups combined. These repeated origins of social parasitism are probably due to the allodapine trait of rearing brood in communal tunnels, a trait that might allow other species to surreptitiously lay additional eggs without them being detected.
Several studies have shown that allodapine bees first evolved in Africa and then spread to Madagascar, Asia and Australia. The earliest dispersal from Africa to Australia occurred about 30 million years ago and did not appear to involve a route via Asia, leading to a biogeographical puzzle because of the expanse of the Indian Ocean separating Australia from Africa. [18] The most likely routes involved were now-submerged island stepping stones across the Indian Ocean, or dispersal from Africa to Antarctica and then overland dispersal from Antarctica to Australia when the two continents were still connected (ref). Both of these scenarios are problematic, but have been suggested for other animal and plant species. [19]
Recent studies are marked by the number of species they have involved that have not been formally described (refs). This suggests that there is a large amount of allodapine diversity that is not covered by formal scientific taxonomy. Conservation concerns centre on two regions: (i) large-scale habitat loss in Madagascar poses a major threat to that island's unique bee fauna, including allodapine bees, many of which are still to be scientifically described; [20] and (ii) the Australian region is likely to contain many undescribed socially parasitic species [21] which are threatened because of their very small populations sizes. Conservation threats to allodapine bees in Asia have not been studied.
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists.
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamily Apoidea. They are currently considered a clade, called Anthophila. There are over 20,000 known species of bees in seven recognized biological families. Some species – including honey bees, bumblebees, and stingless bees – live socially in colonies while most species (>90%) – including mason bees, carpenter bees, leafcutter bees, and sweat bees – are solitary.
Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. This ovipositor is often modified into a stinger. The young develop through holometabolism — that is, they have a wormlike larval stage and an inactive pupal stage before they reach adulthood.
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in eusocial insects such as ants, wasps, bees, and termites.
Brood parasitism is a subclass of parasitism and phenomenon and behavioural pattern of certain animals, brood parasites, that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own, usually using egg mimicry, with eggs that resemble the host's.
Fish reproductive organs include testes and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused. There may also be a range of secondary organs that increase reproductive fitness. The genital papilla is a small, fleshy tube behind the anus in some fishes, from which the sperm or eggs are released; the sex of a fish can often be determined by the shape of its papilla.
Junonia coenia, known as the common buckeye or buckeye, is a butterfly in the family Nymphalidae. Its range covers much of North America and some of Central America, including most of the eastern half of the US, the lower to middle Midwest, the Southwest, southern Canada, and Mexico. Its habitat is open areas with low vegetation and some bare ground. Its original ancestry has been traced to Africa, which then experiences divergence in Asia. The species Junonia grisea, the gray buckeye, is found west of the Rocky Mountains and was formerly a subspecies of Junonia coenia.
Bombus terrestris, the buff-tailed bumblebee or large earth bumblebee, is one of the most numerous bumblebee species in Europe. It is one of the main species used in greenhouse pollination, and so can be found in many countries and areas where it is not native, such as Tasmania. Moreover, it is a eusocial insect with an overlap of generations, a division of labour, and cooperative brood care. The queen is monogamous which means she mates with only one male. B. terrestris workers learn flower colours and forage efficiently.
Hylaeus is a large and diverse cosmopolitan genus within the bee family Colletidae. This genus is also known as the yellow-faced bees or masked bees. This genus is the only truly globally distributed colletid, occurring on all continents except Antarctica.
Apis andreniformis, or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia.
Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers. Cooperative breeding encompasses a wide variety of group structures, from a breeding pair with helpers that are offspring from a previous season, to groups with multiple breeding males and females (polygynandry) and helpers that are the adult offspring of some but not all of the breeders in the group, to groups in which helpers sometimes achieve co-breeding status by producing their own offspring as part of the group's brood. Cooperative breeding occurs across taxonomic groups including birds, mammals, fish, and insects.
Andrena is a genus of bees in the family Andrenidae. With over 1,500 species, it is one of the largest genera of animals. It is a strongly monophyletic group that is difficult to split into more manageable divisions; currently, Andrena is organized into 104 subgenera. It is nearly worldwide in distribution, with the notable exceptions of Oceania and South America. Bees in this genus are commonly known as mining bees due to their ground-nesting lifestyle.
The East African lowland honey bee is a subspecies of the western honey bee. It is native to central, southern and eastern Africa, though at the southern extreme it is replaced by the Cape honey bee. This subspecies has been determined to constitute one part of the ancestry of the Africanized bees spreading through North and South America.
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.
Eusociality is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.
Eusociality evolved repeatedly in different orders of animals, notably termites and the Hymenoptera. This 'true sociality' in animals, in which sterile individuals work to further the reproductive success of others, is found in termites, ambrosia beetles, gall-dwelling aphids, thrips, marine sponge-dwelling shrimp, naked mole-rats, and many genera in the insect order Hymenoptera. The fact that eusociality has evolved so often in the Hymenoptera, but remains rare throughout the rest of the animal kingdom, has made its evolution a topic of debate among evolutionary biologists. Eusocial organisms at first appear to behave in stark contrast with simple interpretations of Darwinian evolution: passing on one's genes to the next generation, or fitness, is a central idea in evolutionary biology.
Within the insect order Hymenoptera, the Halictinae are the largest, most diverse, and most recently diverged of the four halictid subfamilies. They comprise over 2400 bee species belonging to the five taxonomic tribes Augochlorini, Thrinchostomini, Caenohalictini, Sphecodini, and Halictini, which some entomologists alternatively organize into the two tribes Augochlorini and Halictini.
Eucera is a genus of bees in the family Apidae, which comprises more than 100 species. These bees are commonly known as long-horned bees due to their characteristically long antennae, especially in males. Eucera species can be found in diverse habitats, including meadows, fields, and urban gardens, primarily in the Palearctic and Nearctic regions, covering parts of Europe, Asia, North Africa, and North America.
Exoneura robusta is a species of the primitively eusocial allodapine bee, belonging to the genus commonly referred to as "reed bees". Their common name derives from their use of the soft pith of dead fern fronds as a nesting material. They are native to southeastern Australia, living in both montane and heathland habitats. E. robusta do not have a fixed pattern of sociality, but rather they are capable of adapting their social strategy to different environments. While typically univoltine, populations living in warmer habitats are capable of producing two broods per season. This leads to the incidence of sibling rearing and eusocial behavior. E. robusta lack strict morphological castes, thus allowing for their plastic social behavior and dominance hierarchies.
Tetragonula is a genus of stingless bees. In 1961, Brazilian bee expert J.S. Moure first proposed the genus name Tetragonula to improve the classification system by dividing the large genus Trigona stingless bees into 9 smaller groups. About 30 stingless bee species formerly placed in the genus Trigona are now placed in the genus Tetragonula. These bees are found in Oceania, in countries such as Australia, Indonesia, New Guinea, Malaysia, Thailand, the Philippines, India, Sri Lanka, and the Solomon Islands. The most recent tabulation of species listed 31 species.