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Encyclopedia of Science: Wasps

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Britannica Article: Social Insects

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Britannica Page on Wasps

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Britannica Page on Termites

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Britannica Page on Bees

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Britannica Page on Ants

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Podcast: Science Friday on Social Insects

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Social Insects Video

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Voyagers: Alien Empire- FULL VIDEO

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Termites Video

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Encyclopedia of Science: Termites

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Encyclopedia of Science: Bee

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Encyclopedia of Science: Ants

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bumble bees bombus hymenoptera apidae lionel stange fasulo

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Most bumble bees are large, social bees which produce annual colonies. Mated queens overwinter in the soil and emerge from hibernation in early spring when they feed on spring flowers and search for a suitable location, such as a former rodent burrow in the soil, to begin their colonies.

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velvet ants mutillidae hymenoptera jeffrey hertz

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Biological data is very scarce and differences obviously exist between genera and species, especially with regards to their life cycles and development, but the following has been reported for most Mutillidae. Females have the difficult job of locating potential hosts.

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Here's David Attenborough, chilling out on a rock in the middle of Africa, with four lumps of plasticine. The smallest one on the far left represents the brain of a bushbaby, a small primate that lives on its own. The next one is the brain of a colobus monkey, which lives in groups of 15 or so. The one after that is a guenon, another monkey; group size: 25. And on the far right: a baboon that lives in groups of 50. "Were you to give a skull to a researcher who works on monkeys, even though they didn't know what kind of monkey it belonged to, they would be able to accurately predict the size of group in which it lived," says Attenborough.

That sequence, from The Life of Mammals, is a wonderful demonstration of the social brain hypothesis-a bold idea, proposed in the 1980s, which suggests that living in groups drove the evolution of large brains. Social animals face mental challenges that solitary animals do not: they have to recognise the other members of their cliques, cope with fluid and shifting alliances, manage conflicts, and manipulate or deceive their peers. So as social groups get bigger, so should brains. This idea has been repeatedly tested and confirmed in many groups of animals, including hoofed mammals, carnivores, primates, and birds.

What about insects? Ants, termites, bees, and wasps, also live in large societies, and many of them have unusually big brains-at least, for insects. But in 2010, Sarah Farris from West Virginia University and Susanne Schulmeister from the American Museum of Natural History showed that in these groups, large brains evolved some 90 million years before big social groups. If anything, they correlated with parasitic body-snatching rather than group-living.

"That got people thinking," says Sean O'Donnell from Drexel University. "In recent years, there's been a growing rumbling, almost subterranean movement arguing that social brain ideas may not apply to the social insects." His new study is the latest addition to that movement.

O'Donnell's team studied potter wasps, which lead solitary lives, and the closely related paper wasps, which live in colonies of varying sizes and complexities. They collected queens and workers from 29 species of these wasps, carefully dissected their brains, and measured the size of their mushroom bodies-a pair of structures in insect brains that control higher mental abilities like learning and memory.

And to their surprise, they found that as the wasp colonies got bigger, their mushroom bodies got smaller. Even within the social paper wasps, the team found that species with distinct queens and workers-a sign of a more complex society-have similarly sized mushroom bodies to those with no such castes.

"The pattern is so clear," says O'Donnell. "Sociality may actually decrease demands on individual cognition rather than increasing it."

"Here we have the first concerted evidence that costly brains aren't needed to allow sociality, when you can do it other ways," says Robin Dunbar, who first proposed the social brain hypothesis. "There are many routes to sociality."

What other routes? O'Donnell notes that insects and (most) mammals build their societies in fundamentally different ways. Large mammal societies typically include individuals who are distantly related or even unrelated. Insect societies, by contrast, are basically gigantic families, where all the members are either queens (which reproduce) or their descendants (which do not). You could view these colonies less as groups of individuals and more as extensions of the queens.

As such, their members don't particularly need to keep track of shifting relationships, or manage conflicts, or manipulate their peers, or any of the other social challenges that, say, a baboon or a human faces. They have less of a need for bigger and more sophisticated brains.

Social insects also benefit from swarm intelligence, where individuals can achieve astonishing feats of behaviour by following incredibly simple rules. They can build living buildings, raise crops, vaccinate themselves, and make decisions about where to live. In some cases, they make decisions in a way that's uncannily similar to neurons-a colony behaves like a giant brain, and in more than a merely metaphorical way. They have a kind of 'distributed cognition', where many of the mental feats that other animals carry out using a single brain happen at the level of the colony.

Entomologist Seirian Sumner from Bristol University says that there are mammals, like meerkats and banded mongooses, which live in simple societies where adults cooperatively raise their young. These are often compared to primitively social insects, like paper wasps. "They share very similar family structures, group sizes and plasticity in behavioural roles," Sumner says. It would be very interesting to see if the brains of these mammals follow the same patterns as those of O'Donnell's wasps.

O'Donnell is all in favour of more studies. He wants to see if the same patterns hold in other insect groups that include both social and solitary species, including bees and cockroaches. And he's intrigued by the naked mole rats-colonial mammals that have queen and worker castes, much like ants and wasps. "If our ideas are correct, we'd expect to see mole rats following a similar pattern to insects," he says.

Reference: O'Donnell, Bulova, DeLeon, Khodak, Miller & Sulger. 2015. Distributed cognition and social brains: reductions in mushroom body investment accompanied the origins of sociality in wasps (Hymenoptera: Vespidae). Proc Roy Soc B. Citation tbc.

PS: The size of brain regions isn't always the best indicator of intelligence. I asked O'Donnell about this, and he stands by his decision to focus on the mushroom bodies. "It's definitely a blunt tool for studying brain evolution," he says. "Brain tissue is metabolically very expensive, and even if it was just filler, tissue weight is a big deal, especially for a flying insect. We expect there to be really strong constrains on the size of the [mushroom bodies]."

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The Distributed Brainpower of Social Insects

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Here's David Attenborough, chilling out on a rock in the middle of Africa, with four lumps of plasticine. The smallest one on the far left represents the brain of a bushbaby, a small primate that lives on its own. The next one is the brain of a colobus monkey, which lives in groups of 15 or so.

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Order Hymenoptera - Ants, Bees, Wasps and Sawflies - BugGuide.Net

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An online resource devoted to North American insects, spiders and their kin, offering identification, images, and information.

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Order Blattodea - Cockroaches and Termites - BugGuide.Net

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An online resource devoted to North American insects, spiders and their kin, offering identification, images, and information.

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Hymenoptera

What do they look like?

This group of insects contains many subgroups of wasps, and and also bees and ants, which evolved from wasp ancestors.

There are many different sizes in this group. Some parasitic wasps are so small they can develop into adults inside the eggs of other insects. Others are large and strong predators, over 3 cm long. All have chewing mouthparts, and the adults have a thin connection between their last two body sections (the abdomen and the thorax). All have 4 clear wings as adults, with the front pair larger than the back pair (queen and male ants have wings too, but only for a short time. Bees and wasps have straight antennae, ants often have a permanent bend in theirs.

Parasitic wasps often have a very large needle-like structure at the end of their abdomen. This isn't a stinger, its an ovipositor. They use it to inject their eggs inside a host insect, and some drill through bark, wood, or plant stems to get to their victims. These wasps mostly cannot sting, and they do not have the black and bright yellow colors that you may already know about.

Other wasps are predators, biting and stinging other insects and spiders. The wasps eat them, or carry their prey to their to feed their young. Wasps that sting are usually brightly colored with black and yellow stripes, sometimes red. This is a warning to potential predators that they can sting.

Where do they live?

This is the second most diverse group of insects, only beetles are more diverse. There are over 200,000 species of wasps and their relatives known around the world, and probably at least that many still unknown to science. There are hundreds of species of wasps, bees, and ants here in Michigan.

Wasps and their kin are found all around the world.

What kind of habitat do they need?

Wasps, Bees and Ants are found in just about every habitat on land, except only the coldest polar regions.

How do they grow?

Wasps and their relatives all have complete metamorphosis. From the egg a larva hatches out. It looks a lot like a short fat white worm, but has a distinct head, and may have six small jointed legs. The larva grows and molts (sheds its whole skin) several times before transforming into a pupa. This resting stage has some of the body parts of an adult, but it can't move or feed. Inside, it is transforming into an adult. Eventually an adult emerges from its pupal skin.

How do they reproduce?

Female wasps, bees, and ants can lay dozens to many thousands of eggs, depending on the species. Unless they are parasites they make a nest, and supply their larvae with food to eat.

Some species in this group form colonies where only one or a few females (called queens) lay eggs, and the other females in the colony do not reproduce. Instead they take care of the queens' offspring. A few times a year, some of the offspring fly away to start new colonies.

Females in this group do a lot of parental care. In most species each female builds her own nest, and collects a food supply for each of her offspring. In some species they work cooperatively to build a nest and collect food, and in some species many females tend their sisters and brothers and don't reproduce themselves.

How long do they live?

Most wasps live less than one year, some workers for just a few months. Queens sometimes live for several years.

How do they behave?

Most wasps are active during daylight hours. They are active creatures, following scents to find their food. As noted in the reproduction section, some species live together in nests of relatives. No wasps migrate: if it regularly gets too cold or dry where they live, they go dormant until conditions improve.

How do they communicate with each other?

These insects depend on chemical communication (taste/smell), but they also use other means. Males and females find each other with scent chemicals called pheromones, and ants and social wasps and bees use chemicals to identify nestmates and send warnings and other information. Parasitic wasps sometimes leave scent marks on the host insects to tell any other parasitic wasps that they've already laid eggs there. Some ants can also make noises and vibrations to communicate. Of the three groups, wasps are the most visual. They often hunt by sight.

What do they eat?

Parasitic wasps get most of their protein from the host insect or spider they eat as larvae. Adult parasitic wasps mostly just drink nectar.

Most non-parasitic wasps are predators and scavengers. They feed on dead animals, or hunt insects and spiders, and use their sting to paralyze their prey. They eat their prey themselves or bring some back to the hive to feed growing larvae. Some make individual cells, and put a supply of paralyzed prey animals in there along with an egg. The larva hatches and there is all the food they need. In general wasps are attracted to sugary foods like fruit or high protein foods like meat.

Some wasp species have larvae that eat plants the same way that caterpillars do.

What eats them and how do they avoid being eaten?

Wasps are best known for their ability to give a painful sting, and lots of species do use their stingers to defend themselves and their nests. They also build their nests in places that are hard for predators to reach (either up high or underground) and many build nests out of hard mud to keep their larvae safe. Some wasps that attack other bees or wasps have especially hard exoskeletons for armor.

What roles do they have in the ecosystem?

Wasps are important predators of other insects. Some species are valuable pollinators, and their relatives the bees are the most important pollinators of all.

Do they cause problems?

Some of these species have a painful sting. The social nesting species can actually be dangerous to people: if someone disturbs their nest, they may sting them so many times that the person's life is endangered. Also, some wasps attack beneficial insects like bees.

How do they interact with us?

In general this group of organisms has much more positive effects than negative. They are important enemies of many insect pests, helping protect our crops. They are also important pollinators, allowing our flowers and vegetables to grow and reproduce.

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BioKIDS - Kids' Inquiry of Diverse Species, Hymenoptera, wasps, bees, and ants: INFORMATION

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Hymenoptera This group of insects contains many subgroups of wasps, and and also bees and ants, which evolved from wasp ancestors. There are many different sizes in this group. Some parasitic wasps are so small they can develop into adults inside the eggs of other insects.

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Honey bee workers clustering on a honeycomb.
Photograph by Waugsberg, used under .

Introduction

The most familiar Hymenoptera are the bees, wasps and ants; insects feared by many people because of their stings. However, these make up only a small proportion of the order. The Hymenoptera also includes Ichneumons and other parasitic "wasps" and Sawflies. Unfortunately, there is no English name to embrace the whole order as there is for beetles, bugs and true flies.

Hymenoptera characteristically have two pairs of wings, a large fore pair and a smaller hind pair. These wings are held together by a series of hooks (called a frenulum) and may appear like a single pair to the naked eye. Hymenoptera also tend to have prominent antennae, generally with nine or more segments and biting mouthparts.

The most conspicuous Hymenoptera are the fast-flying, often black and yellow, bees and wasps. Ants, too, are easily noticed when intruding on our lives. However, many of the smaller and less familiar Hymenoptera can have a jewel-like beauty on close examination and are interesting in a great variety of ways.

In the British Isles, there are at least 6,700 species of Hymenoptera, occurring in all terrestrial habitats. On a world scale are at least 100,000 species, so this is only a brief introduction to the group and deals mainly with the main British groups.

Hymenoptera and Man

Honey

Honey is associated with all manner of positive things; so often so that it is easy to forget that it is regurgitated by insects! A concentrated form of flower nectar stored by certain bees for their young, honey has been sought since earliest times. There is a 9000-year-old cave painting in Spain depicting the gathering of wild honey and beehives are illustrated on friezes in ancient Egyptian tombs. Honey bees are also valued for their beeswax, royal jelly (used to supplement the diet of developing queens) and other products. Find out more about bees and beekeeping.

Pollination

Many winged Hymenoptera visit flowers. Bees do so to collect pollen and nectar for themselves or to provision their developing young, while wasps are often on the lookout for other insects to prey upon. Flowering plants have, in an evolutionary sense, taken great advantage of this by offering nectar to insects most likely to carry their pollen to another flower. With many important crops dependent on insect pollinators, Hymenoptera have a huge role to play in feeding the world's human population. Bumblebees are routinely introduced into commercial greenhouses where pollinators are in short supply, and the contribution pollination makes to the world's economy is tens of billions of dollars per annum.

Pest control

A parasitic wasp, Peristenus digoneutis, laying an egg in a plant bug nymph.
Scott Bauer,

Wasps are a gardener's best friend killing countless garden pests. In fact, Ichneumons and other Parasitica are used as successful biological control agents. For example, the parasitic wasp Habrolepis dalmanni (Encyrtidae) was introduced from North America into New Zealand for the control of Golden Oak Scale insect Asterolecanium variolosum ( Hemiptera:Asterolecaniidae), a serious pest of oak. Ants are also valued in some forests for their role in controlling pests in large numbers.

That said, not all Hymenoptera are quite so welcome. The larvae of Sawflies and Wood Wasps damage crops and ants can encourage aphid blooms by protecting the aphids from predators.

Life Cycles

Life stages

Hymenoptera undergo complete metamorphosis - that is, they have an egg, larva, pupa and adult stage. The larvae are usually soft and white with no legs, looking similar to diptera larvae; the larvae of some Sawflies have small fore and hind legs like caterpillars.

The sexes are separate, but males are rare in several Parasitica, and the sexes are "redefined" a little in the social aculeates. Many Hymenoptera are protandrous, that is the adult male starts to emerge before the female, awaiting the emergence of females to mate with or even mating before or while she is trying to emerge. In contrast, females of some Parasitica will wait for their own male offspring to emerge before promptly mating with them!

Types of life cycle

There are a bewildering variety of types of life cycle among Hymenoptera. These are intimately bound up with the way in which food is provided for the offspring.

In the Symphyta, eggs are laid in plant tissue and simply abandoned. Other plant-feeding Hymenopterans, such as the Gall Wasps (Parasitica), induce plants to grow protective structures ( galls) that shelter the young as they feed. In the Parasitica, and some aculeates, parasitoid life cycles are common. Parasitoid larvae behave like parasites, feeding off the tissues of living animals, but unlike true parasites usually kill the host at the end of their development. Parasitoids are often extremely specialised, attacking only a single species. When this itself is the larva of a parasitoid insect, they may be called hyperparasitoids.

Nest-building is well-developed in the aculeates, with social organisation being a special case. Solitary, as opposed to social, aculeates may build one or more nests, each supplied with food for the larvae. Other Hymenoptera take advantage of the hard work done by others to supply food. This is known as kleptoparasitism, and the species are often called "cuckoo wasps" or "cuckoo bees". Eggs are laid on another species' store of food, which is eaten by the developing larva, usually killing the rightful heir in the process. There are even social kleptoparasites, in which a whole colony of bees, wasps or ants is taken over from within by another, often closely related, species.

Social Organisation

The phenomenon of sociality among insects is best developed among the aculeate Hymenoptera (it also occurs in termites and a single mammal, the naked mole rat).

Social animals, in this sense, are those in which individuals have evolved different reproductive roles, with at least two types ( castes) of female (a fertile queen and sterile workers) that co-operate in raising the young.

A social colony is like a single organism in that it has a single set of genes to be passed on at reproduction, in the form of a new queen or queens. In the most advanced cases, seen in ants, some bees, and some wasps, the different castes are strikingly different in appearance from one another.

It is in these groups that many fascinating co-operative behaviours have evolved, for example, the paper-making habits of wasps, the waggle-dance communication of honeybees, and the seemingly suicidal defence of colonies by workers with no reproductive interests other than that of the colony as a whole.

The apparent selflessness of honeybees still inspires admiration, as it did to Shakespeare's Henry V: Creatures that by a rule in nature teach / The act of order to a peopled kingdom. It is only since Shakespeare's time that it became widely known that bees have a queen and not a king; and more recently still that the complexities of hymenopteran societies have begun to be fully appreciated.

Diversity and Classification

The Hymenoptera are divided into two suborders, the Symphyta (Sawflies and their relatives) and the Apocrita (species with a thin "waist" including all the wasps, ants and bees). Within the Apocrita, there are two groups, the Parasitica (the most diverse of all Hymenopteran groups) and the Aculeata, species that have modified their egg laying tube ( ovipositor) into a sting. The list below summaries the groups that are dealt with in this introduction to the Hymenoptera, although there is not space here to go into the full classification and evolutionary history of the group.

Essential reading from the Amateur Entomologists' Society

Resources

Related links: Bees, Wasps and Ants (Order: Hymenoptera)

Books

Hymenoptera in general

Gauld, I.D., Bolton, B., Huddleston, T., Fitton, M.G., Shaw, M.R., Noyes, J.S. , Day, M.C., Else, G.R, Fergusson, N.D.M. & Ward, S.L. The Hymenoptera. British Museum (Natural History) Oxford University Press, 1988.

Bethyloids

Perkins, J.F. Hymenoptera Bethyloidea (excluding Chrysidae). Handbooks for the Identification of British Insects, 6, Part 3(a). London: Royal Entomological Society, 1976.

Ants

Bolton, B. & Collingwood, C.A. Hymenoptera, Formicidae. Handbooks for the Identification of British Insects, 6, Part 3c. London: Royal Entomological Society, 1975.

Skinner, G.J. & Allen, G.W. Ants. Naturalists Handbooks 24. Slough: Richmond Publishing Co Ltd, 1966.

Cuckoo Wasps

Morgan, D. Cuckoo wasps (Hymenoptera, Chrysididae). Handbooks for the Identification of British Insects, 6, Part 5. London: Royal Entomological Society, 1984.

Spider Wasps

Day, M.C. Spider wasps (Hymenoptera:Pompilidae). Handbooks for the Identification of British Insects, 6, Part 4. London: Royal Entomological Society, 1988.

Social Wasps

Else, G.R. Identification. Social wasps. British Wildlife, 5, 304-311, 1994.

Potter and Mason Wasps

Archer, M. E. The British Potter and Mason Wasps. A Handbook. Published by the author, 2003, 2nd ed.

Solitary Wasps

Lomholdt, O. The Sphecidae of Fennoscandia and Demark. 2nd edition. Fauna Entomologica Scandinavica, 4. Leiden: Brill, 1984.

Richard, O. W. Scolioidea, Vespoidea and Sphecoidea (Hymentoptera, Aculeata). Handbooks for the identification of British Insects, 6. Part 3(b). London: Royal Entomological Society, 1980.

Yeo, P.F. & Corbet, S.A. Solitary wasps. Naturalists Handbooks 3. The Richmond Publishing Co. Ltd, 1995, 2nd ed.

Bees

Else, G.R. In prep. Handbook of the bees of the British Isles.

Perkins, R.C.L. The British species of Andrena and Nomada. Transactions of the Entomological Society of London, 1919, 218-319.

Perkins, R.C.L. The British species of Halictus and Sphecodes. Entomologist's Monthly Magazine, 58, 46-52, 94-101, 167-174, 1922.

Perkins, R.C.L. Notes on some aculeate Hymenoptera, with corrections of errors. Entomologist's Monthly Magazine, 67, 20, 1931.

Saunders, E. The Hymenoptera Aculeata of the British Isles. London: Reeve & Co.Ltd, 1896.

Bumblebees

Edwards, M. & Jenner, M. Field Guide to the Bumblebees of Great Britain and Ireland. Published by Ocelli Ltd.

Prys-Jones, O. & Corbet, S.A. Bumblebees. Naturalists Handbook 6. The Richmond Publishing Co. Ltd, 1991, Second edition, revised.

Sawflies

Benson, R.B. Hymenoptera: Symphyta. Handbooks for the Identification of British Insects, 6, Part 2(a). London: Royal Entomological Society, 1951.

Benson, R.B. Hymenoptera: Symphyta. Handbooks for the Identification of British Insects, 6, Part 2(b). London: Royal Entomological Society, 1952.

Benson, R.B. Hymenoptera: Symphyta. Handbooks for the Identification of British Insects, 6, Part 2(c). London: Royal Entomological Society, 1958.

Wright, A. British Sawflies a key to the adults of genera occurring in Britain. Field Studies Council, 1990.

Back to Insect Orders.

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Bees, Wasps and Ants (Order: Hymenoptera)

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Honey bee workers clustering on a honeycomb.Photograph by Waugsberg, used under . The most familiar Hymenoptera are the bees, wasps and ants; insects feared by many people because of their stings. However, these make up only a small proportion of the order. The Hymenoptera also includes Ichneumons and other parasitic "wasps" and Sawflies.

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Termites (Order: Isoptera) - Amateur Entomologists' Society (AES)

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Information on Termites (Order: Isoptera). Recent research has shown that termites are actually a lineage of cockroaches.

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Amateur Entomologists Society
BioKids: Insects
Bug Guide
Classification of Insect Orders
Butterflies and Moths
EOL: Insect Orders
Florida 4H Bug Club
Invertebrate Zoology
Kentucky Critter Files
Orkin: Insects
Penn State: Entomology
Seattle Bug Safari: Meet the Insects
Texas Insects