Allodapini

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Allodapini
Allodapula variegata, Pretoria NBT, a.jpg
Allodapula sp. in South Africa
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Apidae
Subfamily: Xylocopinae
Tribe: Allodapini
Cockerell, 1902
Genera

Allodape
Allodapula
Braunsapis
Brevineura
Compsomelissa
Effractapis
Eucondylops
Exoneura
Exoneurella
Exoneuridia
Halterapis
Hasinamelissa
Inquilina
Macrogalea
Nasutapis

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]

Social evolution

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]

Sex allocation

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]

Social parasitism

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.

Historical biogeography

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]

Conservation issues and biodiversity

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.

Related Research Articles

Bee Clade of insects

Bees are flying insects closely related to wasps and ants, known for their role 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 presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families. Some species – including honey bees, bumblebees, and stingless bees – live socially in colonies while some species – including mason bees, carpenter bees, leafcutter bees, and sweat bees – are solitary.

Hymenoptera Order of insects comprising sawflies, wasps, bees, and ants

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.

Parasitoid Organism that lives with host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

Sympatric speciation A process through which new species evolve from a single ancestral species while inhabiting the same geographic region

Sympatric speciation is the evolution of a new species from a surviving ancestral species while both continue to inhabit the same geographic region. In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organisms whose ranges overlap so that they occur together at least in some places. If these organisms are closely related, such a distribution may be the result of sympatric speciation. Etymologically, sympatry is derived from the Greek roots συν ("together") and πατρ

Brood parasite Organism that relies on others to raise its young

Brood parasites are organisms 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, using brood mimicry, for example by having eggs that resemble the host's.

Xylocopinae Subfamily of bees

The subfamily Xylocopinae occurs worldwide, and includes the large carpenter bees, the small carpenter bees, the allodapine bees, and the relictual genus Manuelia.

Fish reproduction The reproductive physiology of fishes

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 often can be determined by the shape of its papilla.

<i>Nomada</i> Genus of bees

With over 850 species, the genus Nomada is one of the largest genera in the family Apidae, and the largest genus of kleptoparasitic "cuckoo bees." Kleptoparasitic bees are so named because they enter the nests of a host and lay eggs there, stealing resources that the host has already collected. The name "Nomada" is derived from the Greek word nomas (νομάς), meaning "roaming" or "wandering."

Reproductive suppression

Reproductive suppression involves the prevention or inhibition of reproduction in otherwise healthy adult individuals. It includes delayed sexual maturation (puberty) or inhibition of sexual receptivity, facultatively increased interbirth interval through delayed or inhibited ovulation or spontaneous or induced abortion, abandonment of immature and dependent offspring, mate guarding, selective destruction and worker policing of eggs in some eusocial insects or cooperatively breeding birds, and infanticide, and infanticide in carnivores) of the offspring of subordinate females either by directly killing by dominant females or males in mammals or indirectly through the withholding of assistance with infant care in marmosets and some carnivores. The Reproductive Suppression Model argues that "females can optimize their lifetime reproductive success by suppressing reproduction when future [physical or social] conditions for the survival of offspring are likely to be greatly improved over present ones". When intragroup competition is high it may be beneficial to suppress the reproduction of others, and for subordinate females to suppress their own reproduction until a later time when social competition is reduced. This leads to reproductive skew within a social group, with some individuals having more offspring than others. The cost of reproductive suppression to the individual is lowest at the earliest stages of a reproductive event and reproductive suppression is often easiest to induce at the pre-ovulatory or earliest stages of pregnancy in mammals, and greatest after a birth. Therefore, neuroendocrine cues for assessing reproductive success should evolve to be reliable at early stages in the ovulatory cycle. Reproductive suppression occurs in its most extreme form in eusocial insects such as termites, hornets and bees and the mammalian naked mole rat which depend on a complex division of labor within the group for survival and in which specific genes, epigenetics and other factors are known to determine whether individuals will permanently be unable to breed or able to reach reproductive maturity under particular social conditions, and cooperatively breeding fish, birds and mammals in which a breeding pair depends on helpers whose reproduction is suppressed for the survival of their own offspring. In eusocial and cooperatively breeding animals most non-reproducing helpers engage in kin selection, enhancing their own inclusive fitness by ensuring the survival of offspring they are closely related to. Wolf packs suppress subordinate breeding.

Neriidae Family of flies

The Neriidae are a family of true flies (Diptera) closely related to the Micropezidae. Some species are known as cactus flies, while others have been called banana stalk flies and the family was earlier treated as subfamily of the Micropezidae which are often called stilt-legged flies. Neriids differ from micropezids in having no significant reduction of the fore legs. Neriids breed in rotting vegetation, such as decaying tree bark or rotting fruit. About 100 species are placed in 19 genera. Neriidae are found mainly in tropical regions, but two North American genera occur, each with one species, and one species of Telostylinus occurs in temperate regions of eastern Australia.

A social spider is a spider species whose individuals form relatively long-lasting aggregations. Whereas most spiders are solitary and even aggressive toward other members of their own species, some hundreds of species in several families show a tendency to live in groups, often referred to as colonies.

Eusociality Highest level of animal sociality a species can attain

Eusociality, the highest level of organization of sociality, 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 at least one behavior characteristic of individuals in another caste.

Evolution of eusociality Origins of cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups.

Eusociality evolved repeatedly in different orders of animals, particularly 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 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.

Halictinae Subfamily of bees

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.

<i>Eucera</i> Genus of bees

Eucera is a genus of bees in the family Apidae, subfamily Apinae, and tribe Eucerini – the long-horned bees.

Sexual selection in amphibians

Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females known to make their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.

Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. The inbreeding avoidance hypothesis posits that certain mechanisms develop within a species, or within a given population of a species, as a result of assortative mating, natural and sexual selection in order to prevent breeding among related individuals in that species or population. Although inbreeding may impose certain evolutionary costs, inbreeding avoidance, which limits the number of potential mates for a given individual, can inflict opportunity costs. Therefore, a balance exists between inbreeding and inbreeding avoidance. This balance determines whether inbreeding mechanisms develop and the specific nature of said mechanisms.

<i>Exoneura robusta</i> Species of bee

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.

<i>Tetragonula hockingsi</i> Species of bee

Tetragonula hockingsi is a small stingless bee native to Australia. It is found primarily in the Northern Territory and in northern Queensland. The colonies can get quite large, with up to 10,000 workers and a single queen. Workers of Tetragonula hockingsi have been observed in fatal fights with other Tetragonula species, where the worker bees risk their lives for the potential benefit of scarce resources.

<i>Tetragonula</i> Genus of bees

Tetragonula is a genus of stingless bees. In 1961, Brazilian bee expert, Professor 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.

References

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  19. Barker, N.P.; Weston, P.H.; Rutschmann, F.R. & Sauquet, H. (2007). "Molecular dating of the 'Gondwanan' plant family Proteaceae is only partially congruent with the timing of the break-up of Gondwana". Journal of Biogeography. 34 (12): 2012–2027. doi:10.1111/j.1365-2699.2007.01749.x.
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  21. Smith, J.A.; Schwarz, M.P. (2006). "New species and unexpected diversity of socially parasitic bees in the genus Inquilina Michener". Insect Science. 16 (4): 343–350. doi:10.1111/j.1744-7917.2009.01266.x.