Mass provisioning is a form of parental investment [1] in which an adult insect, most commonly a hymenopteran such as a bee or wasp, stocks all the food for each of her offspring in a small chamber (a "cell") before she lays the egg. This behavior is common in both solitary and eusocial bees, though essentially absent in eusocial wasps.
In bees, stored provisions typically consist of masses of mixed pollen and nectar, though a few species store floral oils. [2] In a few cases, such as stingless bees and some sweat bees, the number of cells in a single nest can number in the hundreds to thousands, but more typically a nest contains either a single cell, or a small number (fewer than 10). [3]
In predatory wasps, the food is typically in the form of paralyzed or dead prey items; after digging the nest they quickly catch one or a few prey animals, bring them to the nest and lay eggs on them, seal the nest and leave. Some wasp lineages (e.g. Crabronidae) show variation, with some species practicing mass provisioning, while related species may bring back prey after the egg has hatched, and then seal the nest (such "delayed provisioning" is considered to be a stage in the evolution of progressive provisioning [4] and thus of parental care in insects [5] ), or re-open the nest and add more prey items as the larva grows, which is genuine progressive provisioning. [6]
In 1958, Howard E. Evans published a study of the nesting behaviour of Sphecini digger wasps, showing a range of ways of stocking their nests. In Prionyx , several Nearctic and Palaearctic species catch a grasshopper, and then dig a nest for it, so there is one prey per nest. The nest consists of a single cell, and the egg is laid touching the coxa of a hind leg. In contrast, a Neotropical species, P. spinolae , digs the nest first, creating multiple cells, and stocks each cell with 5–10 grasshoppers; the egg is laid on the underside of the thorax. [7]
No eusocial wasp species carries out mass provisioning in the strict sense, though the vespid wasp genus Brachygastra stores provisions of honey in its nests; the honey is used to supplement larval feeding (larvae are still fed masticated prey items, for protein), and also eaten by adults. [8]
The best-known examples from outside the Hymenoptera are dung beetles, which typically provision with either leaves or dung. Once the provisions are in place and the egg is laid, the cell is sealed, to protect the developing brood. [9] [5]
While mass provisioning is typical of some eusocial lineages, such as some sweat bees and all stingless bees, many other eusocial insects, such as ants and honey bees, instead practise progressive provisioning, where the larvae are fed directly and continually during their development; as such, both highly eusocial and primitively eusocial lineages can perform either type of provisioning. [4]
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.
Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees, carpenter bees, orchid bees, cuckoo bees, and a number of other less widely known groups. Many are valuable pollinators in natural habitats and for agricultural crops.
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.
The Polistinae is a subfamily of eusocial wasps belonging to the family Vespidae. They are closely related to the wasps and true hornets of the subfamily Vespinae, containing four tribes. With about 1,100 species total, it is the second-most diverse subfamily within the Vespidae, and while most species are tropical or subtropical, they include some of the most frequently encountered large wasps in temperate regions.
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.
The gyne is the primary reproductive female caste of social insects. Gynes are those destined to become queens, whereas female workers are typically barren and cannot become queens. Having a queen is what makes a "queenright" hive, nest, or colony of eusocial insects. A colony with multiple queens is said to be a polygyne form, whereas one with only one is a monogyne form.
Brachygastra mellifica, commonly known as the Mexican honey wasp, is a neotropical social wasp. It can be found in North America. B. mellifica is one of few wasp species that produces honey. It is also considered a delicacy in some cultures in Mexico. This wasp species is of use to humans because it can be used to control pest species and to pollinate avocados.
Philanthus gibbosus, which is commonly referred to as a beewolf due to its predation practices, is a species of bee-hunting wasp and is the most common and widespread member of the genus in North America. P. gibbosus is of the order Hymenoptera and the genus Philanthus. It is native to the Midwestern United States and the western Appalachians. P. gibbosus are often observed to visit flowers and other plants in search of insect prey to feed their young. The prey that P. gibbosus catches is then coated in a layer of pollen and fed to the young wasps.
Progressive provisioning is a term used in entomology to refer to a form of parental behavior in which an adult feeds its larvae directly after they have hatched, feeding each larva repeatedly until it has completed development. The food is typically in the form of masticated or immobilized prey items, or regurgitated nectar mixed with pollen ; only rarely are other sorts of food resources used. While this sort of direct and repetitive feeding of offspring is extremely common in groups such as birds and mammals, it is far less common among insects, with the exception of eusocial insects. Accordingly, progressive provisioning is universal among ants, and widespread among the social bees and wasps. Certain nonsocial wasps also rear their young with this type of feeding. Young termites are able to feed themselves, and therefore do not demonstrate any form of provisioning.
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, 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. Eusocial colonies can be viewed as superorganisms.
Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes metricus can be found attached to the sides of buildings, trees, and shrubbery.
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.
Ammophila sabulosa, the red-banded sand wasp, is a species of the subfamily Ammophilinae of the solitary hunting wasp family Sphecidae, also called digger wasps. Found across Eurasia, the parasitoid wasp is notable for the mass provisioning behaviour of the females, hunting caterpillars mainly on sunny days, paralysing them with a sting, and burying them in a burrow with a single egg. The species is also remarkable for the extent to which females parasitise their own species, either stealing prey from nests of other females to provision their own nests, or in brood parasitism, removing the other female's egg and laying one of her own instead.
Tetragonisca angustula is a small eusocial stingless bee found in México, Central and South America. It is known by a variety of names in different regions. A subspecies, Tetragonisca angustula fiebrigi, occupies different areas in South America and has a slightly different coloration.
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.
Melipona bicolorLepeletier, 1836, commonly known as Guaraipo or Guarupu, is a eusocial bee found primarily in South America. It is an inhabitant of the Araucaria Forest and the Atlantic Rainforest, and is most commonly found from South to East Brazil, Bolivia, Argentina, and Paraguay. It prefers to nest close to the soil, in hollowed trunks or roots of trees. M. bicolor is a member of the tribe Meliponini, and is therefore a stingless bee. This species is unique among the stingless bees species because it is polygynous, which is rare for eusocial bees.
Melipona beecheii is a species of eusocial stingless bee. It is native to Central America from the Yucatán Peninsula in the north to Costa Rica in the south. M. beecheii was cultivated in the Yucatán Peninsula starting in the pre-Columbian era by the ancient Maya civilization. The Mayan name for M. beecheii is xunan kab, which translates roughly to "regal lady bee". M. beecheii serves as the subject of various Mayan religious ceremonies.
Scaptotrigona postica is a species of stingless bee that lives mainly in Brazil. It is a eusocial bee in the tribe Meliponini. S. postica is one of 25 species in the genus Scaptotrigona and is a critical pollinator of the tropical rain forests of Brazil. They construct their nests in hollowed sections of tree trunks, allowing for effective guarding at the nest entrance. This species shows colony structure similar to most members of the Meliponini tribe with three roles within the colony: queen, worker, and male. S. postica individuals have different forms of communication from cuticular hydrocarbons to pheromones and scent trails. Communication is especially useful during worker foraging for nectar and pollen through the Brazilian tropical rain forests. S. postica is a very important pollinator of the Brazilian tropical rain forests and is widely appreciated for its honey. Stingless bees account for approximately 30% of all pollination of the Brazilian Caatinga and Pantanal ecosystems and up to 90% of the pollination for many species of the Brazilian Atlantic Forest and the Amazon.
Melipona quadrifasciata is a species of eusocial, stingless bee of the order Hymenoptera. It is native to the southeastern coastal states of Brazil, where it is more commonly known as mandaçaia, which means "beautiful guard," as there is always a bee at the narrow entrance of the nest. M. quadrifasciata constructs mud hives in the hollows of trees to create thin passages that only allow one bee to pass at a time. Because they are stingless bees, M. quadrifasciata is often used as pollinators in greenhouses, outperforming honey bees in efficiency and leading to overall larger yields of fruits that were heavier, larger, and contained more seeds.