Megalopta

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Megalopta
Megalopta sp. (36581769543).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Halictidae
Subfamily: Halictinae
Genus: Megalopta
Smith, 1853

Megalopta is a widespread neotropical genus of bees in the tribe Augochlorini in family Halictidae, known as the sweat bees. [1] They are the largest of the five nocturnal genera in Augochlorini. Most have pale integumentary pigmentation, and all have large ocelli, most likely a feature of their nocturnal behavior. [2] They live in tropical Central America and the entirety of South America. The subgenus Noctoraptor is cleptoparasitic. [3] They are not known from the fossil record. [4]

Contents

Megalopta was first described by Frederick Smith in 1853. [5] The type species is Megalopta idalia, now known as Megalopta amoena . [5] [6] Most studies done on Megalopta are focused on M. genalis.

General description

Megalopta are up to 2 cm long. They have large ocelli and compound eyes, used for nocturnal foraging. [7] They have a yellow/brown abdomen with dark brown banding and metallic green/yellow to bronze thorax and head. [1]

Nocturnal behavior and vision

All species of Megalopta are believed to be nocturnal or crepuscular. [4] In general, Megalopta exhibit bimodal foraging patterns and are active for about 90 minutes after the sun sets and before the sun rises, [8] regardless of food availability. [9] Males and females are active at slightly different times and the females usually have higher overall rates of activity. [8] This is hypothesized to be a niche shift to avoid competition from diurnal bees, or perhaps to avoid predation [9]

Insects generally have a pair of compound eyes accompanied by a trio of single-lensed ocelli on the top of their head. [10] While most nocturnal insects have superposition compound eyes, all hymenoptera have apposition eyes that are less sensitive to light than superposition eyes. Megalopta's apposition eyes are thirty times more sensitive to light than diurnal bees, but even this does not explain the accuracy of their vision. [11] They may use their ocelli to supplement their night vision as well. Crepuscular bees have larger ocelli than diurnal bees, and nocturnal bees have the largest ocelli of all. [12] Ocelli are not used for precise visual information, but can be used for other uses of visual information. Megalopta ocelli are highly adapted for sensitivity. They have large ocellar lenses and the percent area of the retina covered by the rhabdom is five times higher than diurnal bees. [10] However, the adaptation to sensitivity most likely came at the cost of temporal resolution. One theory is they use neural spatial summation strategies to enhance their temporal resolution. [13] Their rhabdom are elongated, with microvilli aligned in the same direction which potentially allows them to see the direction of polarized light. [10] Megalopta ocelli have a dorsal rim, where the photoreceptors are sensitive to polarized light, such as from the rising or setting sun. Sunset and sunrise create unique polarized light patterns in the sky, and Megalopta may be using these light patterns to navigate. [14] Despite having the superficial anatomy for polarized light directionality detection, the rhabdom are "worm-shaped" and arranged radially, which negates this ability. [10] Meglopta also lack pigments in their ocelli that detect short-wavelength light, perhaps a result of their jungle habitat. [10]

Nesting and the female hierarchy

Megalopta nest in dead branches, sticks, and vines. Their nests have a distinctive collar around the entrance made of chewed wood and consist of a large hollow tunnel and several cells. [4] The internal cells are made of the same chewed wood as the entrance. Megalopta are facultatively eusocial. [2] Their nests range from single female nests up to eleven female nests, but social nests usually contain 2-4 females. [15] As the dry season progresses, single female nests gradually shift into multiple female nests, a result of females hatching and remaining in the nest. [4] Usually, at least one female offspring stays in the nest and does not reproduce or grow ovaries. She instead maintains the nest and forages while the foundress continues to reproduce. [16] Eusocial communities have an overlap of generations of nest-mates, meaning that two or more broods must be laid and hatch during the mating season. [17] Eggs are usually laid during the dry season and hatch after 35 days. [4] [17] Eggs laid in a drier dry season usually produce numerically more, larger individuals that are involved in reproduction, as opposed to a comparatively wet dry season, which produces fewer, smaller workers. [17] The association between reproduction and the dry season is likely a result of floral availability. [1] The benefits of solitary versus eusocial lifestyles change with the seasons. In the dry season, eusocial lifestyles provide productivity benefits while in the wet season, the benefits are typically insurance-based. [18]

In a multi-female hive, there is typically one dominant female and the rest are considered supernumerary. [17] The dominant female usually is macrocephalic and the oldest female in the hive. [3] The dominant female is analogous to the queen in honeybee hives, and can also be called the queen or the foundress. The supernumerary females fall into two roles: foragers and in-nest. The in-nest females generally stay in the nest and guard the entrance, as well as hygiene-oriented tasks. [15] Halictinae are mass-provisional, meaning they cache the food necessary for larval growth before the eggs are laid and do not interact with larvae during development. [16] However, experimentally, foundresses have displayed the ability to identify issues within a sealed cell. The foundresses opened the cell to assess the condition of the larvae and consequently either found something wrong, aborted the larvae, and reused the cell, or found nothing wrong and resealed the cell. The larvae in the resealed cells developed normally. [16] Another parental behavior performed by foundresses is hygienic-based: the removal of faeces from cells that produced healthy adults. [16] These deviations from mass-provisional behaviors are thought to be a precursor to eusociality. [16]

Gynandromorphy

Gynandromorphy is the presence of both female and male characteristics in a single organism. Two species of Megalopta have exhibited gynandromorphy: Megalopta amoena and the heavily studied Megalopta genalis. Both specimens had a bilateral split with male characteristics on the left and female characteristics on the right. In comparison to male or female individuals who have bimodal foraging periods, the gynandromorph's activity was shifted significantly earlier in the day. [8] It is important to note that this study was based on a single specimen and generalizations about gynandromorphy in Megalopta cannot be made. Furthermore, this mutation is extremely rare, with only two specimens having been found in the entire genus.

Interactions with other species

Macrosiagon gracilis is a parasite of M. genalis and M.ecuadoria. [19] Megalopta have lower rates of brood parasitism than other solitary bees. Significant brood parasites include Lophostigma cincta , a mutilid wasp, and cleptoparasitic Megalopta species. [9] In many dead nests, slits are found in the stick and the cells are destroyed. This is thought to be the work of the silky anteater, Cyclopes didactylus , but is unconfirmed. [4] Females also often have large numbers of nematodes in their metasomal glands. [4] The larvae of a parasitic fly, Fiebrigella sp., consume the pollen stores in Megalopta cells, causing the larvae inhabiting those cells to have a smaller body size. [15] Parkia velutina is pollinated by Megalopta [7]

Cleptoparasitism

Cogener cleptoparasitism has been observed in the subgenus Noctoraptor. This means that these bees parasitize other Megalopta species. M. byroni parasitizes M. genalis and M. ecuadoria, but most likely parasitizes other species [4] . In the case of Noctoraptor, cleptoparasitism takes the form of brood parasitism. Noctoraptor do not have the physiology to build nests or cells, making them obligate parasites that invade the nests of the other Megalopta to lay eggs.

Species and subgenera

There is considerable discourse on the phylogeny within Megalopta. One school of thought states Megalopta has two subgenera: Megalopta and Noctoraptor. [2] The subgenus Noctoraptor was described in 1997, with the type species of M. byroni. [20] M. byroni was the first known nocturnal parasitic bee. [20] Noctoraptor can be differentiated from other Megalopta by anatomical differences that are linked to parasitism, including a reduced scopa, large scythe-shaped mandibles, and the lack of a basitibial plate. [20] In newer literature, instead of being classified into the two subgenera, the divisions are made between five species groups: Aegis, Amoena, Yanomami, Byroni, and Sodalis. These groups are defined by similar physical characteristics. The Byroni species group contains the species that would be in Noctoraptor. [6] These species groups have been determined for Megalopta in Brazil, so the species group listings do not include all known species. [6] The species group system is not widely used.

Regardless of subgenera or species groups, Megalopta is most likely paraphyletic and is grouped with the genus Xenochlora , with the species M. atra being most closely related to Xenochlora over other Megalopta. [21] Xenochlora may be a subgenus of Megalopta. [21] M. atra is considered a highland species, while the rest of the genus is lowland. Megalopta, Megommation , and Megaloptidia form a single clade with a dim-light foraging ancestor. This implies that the Xenochlora clade reverted to diurnal foraging after the adaptations to nocturnal foraging evolved. [21]

The species Megalopta amoena has been known under several different names: Megalopta ecuadoria, Megalopta centralis, Megalopta idalia, Megalopta gibbosa, Megalopta ochrias, Megalopta lecointei, and Megalopta vigilans. [6]

List of species

Species groups [6]

  • Aegis species group
    • Megalopta aegis
    • Megalopta aeneicollis
    • Megalopta nitidicollis
    • Megalopta sulciventris
  • Amoena species group
    • Megalopta amoena
    • Megalopta chaperi
    • Megalopta guimaraesi
    • Megalopta mura
  • Yanomami species group
    • Megalopta piraha
    • Megalopta yanomami
  • Byroni species group
    • Megalopta atlantica
    • Megalopta byroni
    • Megalopta guarani
    • Megalopta karitiana
    • Megalopta mapinguari
    • Megalopta noctifurax
    • Megalopta purpurata
    • Megalopta xavante
  • Sodalis species group
    • Megalopta cuprea
    • Megalopta munduruku
    • Megalopta sodalis

Related Research Articles

<span class="mw-page-title-main">Bee</span> Clade of insects

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.

<span class="mw-page-title-main">Trophallaxis</span> Transfer of food between members of a community through stomodeal or proctodeal means

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.

<span class="mw-page-title-main">Halictidae</span> Family of bees

Halictidae is the second-largest family of bees with nearly 4,500 species. They are commonly called sweat bees, as they are often attracted to perspiration. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every continent except Antarctica. Usually dark-colored and often metallic, halictids are found in various sizes, colors and patterns. Several species are all or partly green and a few are red, purple, or blue. A number of them have yellow markings, especially the males, which commonly have yellow faces, a pattern widespread among the various families of bees. The family is one of many with short tongues and is best distinguished by the arcuate basal vein found on the wing. Females in this family tend to be larger than the males. They are the group for which the term 'eusocial' was first coined by entomologist, Suzanne Batra.

<i>Halictus rubicundus</i> Species of bee

Halictus rubicundus, the orange-legged furrow bee, is a species of sweat bee found throughout the Northern Hemisphere. H. rubicundus entered North America from the Old World during one of two main invasions of Halictus subgenera. These invasions likely occurred via the Bering land bridge at times of low sea level during the Pleistocene epoch.

<i>Lasioglossum malachurum</i> Species of bee

Lasioglossum malachurum, the sharp-collared furrow bee, is a small European halictid bee. This species is obligately eusocial, with queens and workers, though the differences between the castes are not nearly as extreme as in honey bees. Early taxonomists mistakenly assigned the worker females to a different species from the queens. They are small, shiny, mostly black bees with off-white hair bands at the bases of the abdominal segments. L. malachurum is one of the more extensively studied species in the genus Lasioglossum, also known as sweat bees. Researchers have discovered that the eusocial behavior in colonies of L. malachurum varies significantly dependent upon the region of Europe in which each colony is located.

<i>Agapostemon</i> Genus of bees

The genus Agapostemon is a common group of Western Hemisphere sweat bees.

<span class="mw-page-title-main">Eusociality</span> Highest level of animal sociality a species can attain

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.

<span class="mw-page-title-main">Halictinae</span> 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>Lasioglossum zephyrus</i> Species of bee

Lasioglossum zephyrus is a sweat bee of the family Halictidae, found in the U.S. and Canada. It appears in the literature primarily under the misspelling "zephyrum". It is considered a primitively eusocial bee, although it may be facultatively solitary. The species nests in burrows in the soil.

<i>Halictus ligatus</i> Species of bee

Halictus ligatus is a species of sweat bee from the family Halictidae, among the species that mine or burrow into the ground to create their nests. H. ligatus, like Lasioglossum zephyrus, is a primitively eusocial bee species, in which aggression is one of the most influential behaviors for establishing hierarchy within the colony, and H. ligatus exhibits both reproductive division of labor and overlapping generations.

<i>Megalopta genalis</i> Species of bee

Megalopta genalis is a species of the family Halictidae, otherwise known as the sweat bees. The bee is native to Central and South America. Its eyes have anatomical adaptations that make them 27 times more sensitive to light than diurnal bees, giving it the ability to be nocturnal. However, its eyes are not completely different from other diurnal bees, but are still apposition compound eyes. The difference therefore lies purely in adaptations to become nocturnal, increasing the success of foraging and minimizing the danger of doing so from predation. This species has served as a model organism in studies of social behavior and night vision in bees.

<i>Lasioglossum cressonii</i> Species of insect

Lasioglossum cressonii is a species in the sweat bee genus Lasioglossum, family Halictidae. Halictidae exhibit eusocial hierarchy behavior which is interesting given that eusociality in this group is hard to evolve and easy to lose. L. cressonii is found throughout North America. L. cressonii have been shown to be important pollinators for apple trees and many other North American native plants.

<i>Lasioglossum vierecki</i> Species of bee

Lasioglossum vierecki, also known as Dialictus vierecki and Halictus vierecki, is a sand sweat bee and is part of the family Halictidae of the order Hymenoptera. It is found in the eastern half of North America from Minnesota to the New England States down to Georgia and Louisiana and up in Manitoba and Ontario. Commonly found in sandy areas, it pollinates various flowers such as grass-leaved goldenrod and rattlesnake master.

Lasioglossum figueresi, formerly known as Dialictus figueresi, is a solitary sweat bee that is part of the family Halictidae of the order Hymenoptera. Found in Central America, it nests in vertical earthen banks which are normally inhabited by one, though sometimes two or even three, females. Females die before their larvae hatch. It was named after José Figueres Ferrer, a famous Costa Rican patriot, and studies of its behavior are now general models for social behavior studies.

Lasioglossum aeneiventre, also known as Dialictus aeneiventre, is a social sweat bee and is part of the family Halictidae of the order Hymenoptera. Found in Central America, it nests mostly on flat ground though sometimes in vertical banks. It is often compared to L. figueresi.

<i>Lasioglossum leucozonium</i> Species of bee

Lasioglossum leucozonium, also known as Lasioglossum similis, is a widespread solitary sweat bee found in North America, Europe, Asia, and parts of northern Africa. While now a common bee in North America, population genetic analysis has shown that it is actually an introduced species in this region. This population was most likely founded by a single female bee.

<i>Augochlora pura</i> Species of insect

Augochlora pura is a solitary sweat bee found primarily in the Eastern United States. It is known for its bright green color and its tendency to forage on a variety of plants. Inhabiting rotting logs, this bee can produce up to three generations per year. Both males and females have been observed licking sweat from human skin, most likely seeking salt

<i>Augochlorella</i> Genus of bees

Augochlorella is a genus in the bee family Halictidae, commonly called sweat bees. They display metallic coloration, ranging from reddish to gold to bluish green, as is typical for other genera in the tribe Augochlorini.

<i>Augochloropsis</i> Genus of bees

Augochloropsis is a genus of brilliant metallic, often blue-green, sweat bees in the family Halictidae. There are at least 140 described species in Augochloropsis.

<i>Augochlorella aurata</i> Species of insect

Augochlorella aurata is a primitively eusocial species of sweat bee in the family Halictidae. It is one of three species of Augochlorella found east of the Rocky Mountains in North America. The body is a brilliant green metallic color, diffused to varying extents with a copper, red, or yellow color. A. aurata is a generalist pollen feeder and likely an important pollinator for some horticultural crops. A common name is golden green sweat bee.

References

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