Carl Eduard Adolph Gerstaecker (30 August 1828 – 20 June 1895) was a German zoologist , entomologist and professor at the University of Berlin and then the University of Greifswald .
107-491: Sawflies are wasp -like insects that are in the suborder Symphyta within the order Hymenoptera , alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor , which the females use to cut into the plants where they lay their eggs. The name is associated especially with the Tenthredinoidea , by far the largest superfamily in the suborder, with about 7,000 known species; in
214-433: 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. The most commonly known wasps, such as yellowjackets and hornets , are in the family Vespidae and are eusocial , living together in a nest with an egg-laying queen and non-reproducing workers. Eusociality
321-427: A dagger ( † ). Many species of sawfly have retained their ancestral attributes throughout time, specifically their plant-eating habits, wing veins and the unmodified abdomen, where the first two segments appear like the succeeding segments. The absence of the narrow wasp waist distinguishes sawflies from other members of hymenoptera, although some are Batesian mimics with coloration similar to wasps and bees, and
428-479: A sting used for defense or prey capture. Wasps play many ecological roles . Some are predators or pollinators , whether to feed themselves or to provision their nests. Many, notably the cuckoo wasps , are kleptoparasites , laying eggs in the nests of other wasps. Many of the solitary wasps are parasitoidal , meaning they lay eggs on or in other insects (any life stage from egg to adult) and often provision their own nests with such hosts . Unlike true parasites,
535-639: A body length of only 0.139 mm (0.0055 in), and the smallest known flying insect, only 0.15 mm (0.0059 in) long. Wasps have appeared in literature from Classical times , as the eponymous chorus of old men in Aristophanes ' 422 BC comedy The Wasps , and in science fiction from H. G. Wells 's 1904 novel The Food of the Gods and How It Came to Earth , featuring giant wasps with three-inch-long stings. The name 'Wasp' has been used for many warships and other military equipment. The wasps are
642-516: A body length of only 158 micrometres, the smallest known flying insect. There are estimated to be 100,000 species of ichneumonoid wasps in the families Braconidae and Ichneumonidae . These are almost exclusively parasitoids, mostly using other insects as hosts. Another family, the Pompilidae , is a specialist parasitoid of spiders. Some wasps are even parasitoids of parasitoids; the eggs of Euceros are laid beside lepidopteran larvae and
749-415: A broad connection between the abdomen and the thorax . Some sawflies are Batesian mimics of wasps and bees, and the ovipositor can be mistaken for a stinger . Sawflies vary in length, most measuring 2.5 to 20 millimetres ( 3 ⁄ 32 to 25 ⁄ 32 in); the largest known sawfly measured 55 mm ( 2 + 1 ⁄ 4 in). The larvae are caterpillar-like, but can be distinguished by
856-552: A cosmopolitan paraphyletic grouping of hundreds of thousands of species, consisting of the narrow-waisted clade Apocrita without the ants and bees . The Hymenoptera also contain the somewhat wasplike but unwaisted Symphyta , the sawflies. The term wasp is sometimes used more narrowly for members of the Vespidae , which includes several eusocial wasp lineages, such as yellowjackets (the genera Vespula and Dolichovespula ), hornets (genus Vespa ), and members of
963-401: A crop inside their bodies, rather than on body hairs like bees, and pollinate flowers of Penstemon and the water leaf family, Hydrophyllaceae . The Agaonidae ( fig wasps ) are the only pollinators of nearly 1000 species of figs , and thus are crucial to the survival of their host plants. Since the wasps are equally dependent on their fig trees for survival, the coevolved relationship
1070-406: A danger as the wasps may sting if people come close to them. Stings are usually painful rather than dangerous, but in rare cases, people may suffer life-threatening anaphylactic shock . Some species of parasitic wasp, especially in groups such as Aphelinidae , Braconidae , Mymaridae , and Trichogrammatidae , are exploited commercially to provide biological control of insect pests . One of
1177-400: A family or genus of hosts. For example, Iris sawfly larvae, emerging in summer, can quickly defoliate species of Iris including the yellow flag and other freshwater species. Similarly the rose sawflies, Arge pagana and A. ochropus , defoliate rose bushes. The giant woodwasp or horntail, Urocerus gigas , has a long ovipositor, which with its black and yellow colouration make it
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#17327649600901284-489: A football club in Scotland . Wasps have been modelled in jewellery since at least the nineteenth century, when diamond and emerald wasp brooches were made in gold and silver settings. A fashion for wasp waisted female silhouettes with sharply cinched waistlines emphasizing the wearer's hips and bust arose repeatedly in the nineteenth and twentieth centuries. The Ancient Greek playwright Aristophanes wrote
1391-410: A fur-like covering of soft hairs and a special body part for pollen storage ( pollen basket ) as some bees do, pollen does not stick to them well. However it has been shown that even without hairs, several wasp species are able to effectively transport pollen, therefore contributing for potential pollination of several plant species. Pollen wasps in the subfamily Masarinae gather nectar and pollen in
1498-463: A good mimic of a hornet . Despite the alarming appearance, the insect cannot sting. The eggs are laid in the wood of conifers such as Douglas fir , pine, spruce , and larch . The larvae eat tunnels in the wood, causing economic damage. Alternative measures to control sawflies can be taken. Small-scale, mechanical methods include visually confirming larval presence on a plant and subsequently removing them, either by pruning damaged leaves or removing
1605-826: A humid and warm climate. The cladogram is based on Schulmeister 2003. Xyeloidea ( Triassic –present) [REDACTED] Tenthredinoidea [REDACTED] Pamphilioidea [REDACTED] Cephoidea (stem sawflies) [REDACTED] Siricoidea (horntails or wood wasps) [REDACTED] Xiphydrioidea (wood wasps) [REDACTED] Orussoidea ( parasitic wood wasps) [REDACTED] Apocrita ( ants , bees , wasps ) [REDACTED] There are approximately 8,000 species of sawfly in more than 800 genera, although new species continue to be discovered. However, earlier studies indicated that 10,000 species grouped into about 1,000 genera were known. Early phylogenies such as that of Alexandr Rasnitsyn , based on morphology and behaviour, identified nine clades which did not reflect
1712-446: A hundred thousand described species around the world, and a great many more as yet undescribed. For example, almost every one of some 1000 species of tropical fig trees has its own specific fig wasp ( Chalcidoidea ) that has co-evolved with it and pollinates it. Many wasp species are parasitoids; the females deposit eggs on or in a host arthropod on which the larvae then feed. Some larvae start off as parasitoids, but convert at
1819-413: A later stage to consuming the plant tissues that their host is feeding on. In other species, the eggs are laid directly into plant tissues and form galls , which protect the developing larvae from predators, but not necessarily from other parasitic wasps. In some species, the larvae are predatory themselves; the wasp eggs are deposited in clusters of eggs laid by other insects, and these are then consumed by
1926-416: A leaf. Larvae that feed on wood will pupate in the tunnels they have constructed. In one species, the jumping-disc sawfly ( Phyllotoma aceris ) forms a cocoon which can act like a parachute. The larvae live in sycamore trees and do not damage the upper or lower cuticles of leaves that they feed on. When fully developed, they cut small perforations in the upper cuticle to form a circle. After this, they weave
2033-508: A mutualistic relationship with a polydnavirus that weakens the host's immune system and replicates in the oviduct of the female wasp. One family of chalcidoid wasps, the Eucharitidae , has specialized as parasitoids of ants , most species hosted by one genus of ant. Eucharitids are among the few parasitoids that have been able to overcome ants' effective defences against parasitoids. Many species of wasp, including especially
2140-447: A number of chemicals, including maldison , dimethoate , carbaryl , imidacloprid , etc., if removing larvae from trees is not effective enough. Wasp 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
2247-538: A protected environment; this may be the body of a host organism or a cell in a nest, where the larva either eats the provisions left for it or, in social species, is fed by the adults. Such larvae have soft bodies with no limbs, and have a blind gut (presumably so that they do not foul their cell). Adult solitary wasps mainly feed on nectar, but the majority of their time is taken up by foraging for food for their carnivorous young, mostly insects or spiders. Apart from providing food for their larval offspring, no maternal care
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#17327649600902354-566: A silk hammocks within the circle; this silk hammock never touches the lower cuticle. Once inside, the upper-cuticle's disc separates and descends towards the surface with the larvae attaching themselves to the hammock. Once they reach the round, the larvae work their way into a sheltered area by jerking their discs along. The majority of sawfly species produce a single generation per year, but others may only have one generation every two years. Most sawflies are also female, making males rare. Sawflies are major economic pests of forestry . Species in
2461-524: A stinger for both defence and for paralysing prey. In addition to their large compound eyes , wasps have several simple eyes known as ocelli , which are typically arranged in a triangle just forward of the vertex of the head. Wasps possess mandibles adapted for biting and cutting, like those of many other insects, such as grasshoppers , but their other mouthparts are formed into a suctorial proboscis , which enables them to drink nectar. The larvae of wasps resemble maggots , and are adapted for life in
2568-401: A true brood parasite is the paper wasp Polistes sulcifer , which lays its eggs in the nests of other paper wasps (specifically Polistes dominula ), and whose larvae are then fed directly by the host. Sand wasps Ammophila often save time and energy by parasitising the nests of other females of their own species, either kleptoparasitically stealing prey, or as brood parasites, removing
2675-480: A wide variety of prey, mainly other insects (including other Hymenoptera), both larvae and adults. The Pompilidae specialize in catching spiders to provision their nests. Some social wasps are omnivorous, feeding on fallen fruit, nectar, and carrion such as dead insects. Adult male wasps sometimes visit flowers to obtain nectar . Some wasps, such as Polistes fuscatus , commonly return to locations where they previously found prey to forage. In many social species,
2782-432: A wingspan of 92 mm ( 3 + 1 ⁄ 2 in). The smaller species only reach lengths of 2.5 mm ( 3 ⁄ 32 in). Heads of sawflies vary in size, shape and sturdiness, as well as the positions of the eyes and antennae. They are characterised in four head types: open head, maxapontal head, closed head and genapontal head. The open head is simplistic, whereas all the other heads are derived. The head
2889-517: Is Hymenoptera, the Symphyta has long been seen to be paraphyletic. Cladistic methods and molecular phylogenetics are improving the understanding of relationships between the superfamilies, resulting in revisions at the level of superfamily and family. The Symphyta are the most primitive ( basal ) taxa within the Hymenoptera (some going back 250 million years), and one of the taxa within
2996-512: Is a parasitoid of the Atlas beetle Chalcosoma atlas . The female giant ichneumon wasp Megarhyssa macrurus is 12.5 centimetres (5 in) long including its very long but slender ovipositor which is used for boring into wood and inserting eggs. The smallest wasps are solitary parasitoid wasps in the family Mymaridae , including the world's smallest known insect, Dicopomorpha echmepterygis (139 micrometres long) and Kikiki huna with
3103-599: Is a warning colouration because some larvae can secrete irritating fluids from glands located on their undersides. Sawflies are widely distributed throughout the world. The largest family, the Tenthredinidae, with some 5,000 species, are found on all continents except Antarctica, though they are most abundant and diverse in the temperate regions of the northern hemisphere; they are absent from New Zealand and there are few of them in Australia. The next largest family,
3210-459: Is also hypognathous, meaning that the lower mouthparts are directed downwards. When in use, the mouthparts may be directed forwards, but this is only caused when the sawfly swings its entire head forward in a pendulum motion. Unlike most primitive insects, the sutures (rigid joints between two or more hard elements on an organism) and sclerites (hardened body parts) are obsolescent or absent. The clypeus (a sclerite that makes up an insects "face")
3317-605: Is between 205.6 and 221.5 million years of age, making it among the oldest of all sawfly fossils. More Xyelid fossils have been discovered from the Middle Jurassic and the Cretaceous , but the family was less diverse then than during the Mesozoic and Tertiary . The subfamily Xyelinae were plentiful during these time periods, in which Tertiary faunas were dominated by the tribe Xyelini; these are indicative of
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3424-484: Is favoured by the unusual haplodiploid system of sex determination in Hymenoptera, as it makes sisters exceptionally closely related to each other. However, the majority of wasp species are solitary, with each adult female living and breeding independently. Females typically have an ovipositor for laying eggs in or near a food source for the larvae, though in the Aculeata the ovipositor is often modified instead into
3531-413: Is fully mutualistic . Most solitary wasps are parasitoids. As adults, those that do feed typically only take nectar from flowers. Parasitoid wasps are extremely diverse in habits, many laying their eggs in inert stages of their host ( egg or pupa ), sometimes paralysing their prey by injecting it with venom through their ovipositor. They then insert one or more eggs into the host or deposit them upon
3638-463: Is generally considered Russell's best novel. In Stieg Larsson 's book The Girl Who Played with Fire (2006) and its film adaptation , Lisbeth Salander has adopted her kickboxing ringname, "The Wasp", as her hacker handle and has a wasp tattoo on her neck, indicating her high status among hackers, unlike her real world situation, and that like a small but painfully stinging wasp, she could be dangerous. Parasitoidal wasps played an indirect role in
3745-492: Is given. Some wasp species provide food for the young repeatedly during their development ( progressive provisioning ). Others, such as potter wasps (Eumeninae) and sand wasps ( Ammophila , Sphecidae ), repeatedly build nests which they stock with a supply of immobilised prey such as one large caterpillar, laying a single egg in or on its body, and then sealing up the entrance ( mass provisioning ). Predatory and parasitoidal wasps subdue their prey by stinging it. They hunt
3852-627: Is involved in caring for its own offspring (except for such actions as stealing other wasps' prey or laying in other wasp's nests). There are some species of solitary wasp that build communal nests, each insect having its own cell and providing food for its own offspring, but these wasps do not adopt the division of labour and the complex behavioural patterns adopted by eusocial species. Adult solitary wasps spend most of their time in preparing their nests and foraging for food for their young, mostly insects or spiders. Their nesting habits are more diverse than those of social wasps. Many species dig burrows in
3959-430: Is minimal, and fewer than 10 of these species actually cause a significant impact on sawfly populations. Many of these species attack their hosts in the grass or in other parasitoids. Well known and important parasitoids include Braconidae , Eulophidae and Ichneumonidae . Braconid wasps attack sawflies in many regions throughout the world, in which they are ectoparasitoids, meaning that the larvae live and feed outside of
4066-678: Is most frequent in the anal veins. In all sawflies, 2A and 3A tend to fuse with the first anal vein. This occurs in several families including Argidae, Diprionidae and Cimbicidae. The larvae of sawflies are easily mistaken for lepidopteran larvae (caterpillars). However, several morphological differences can distinguish the two: while both larvae share three pairs of thoracic legs and an apical pair of abdominal prolegs , lepidopteran caterpillars have four pairs of prolegs on abdominal segments 3–6 while sawfly larvae have five pairs of prolegs located on abdominal segments 2–6; crochets are present on lepidopteran larvae, whereas on sawfly larvae they are not;
4173-430: Is not always the case; Monterey pine sawfly ( Itycorsia ) larvae are solitary web-spinners that feed on Monterey pine trees inside a silken web. The adults feed on pollen and nectar. Sawflies are eaten by a wide variety of predators. While many birds find the larvae distasteful, some such as the currawong ( Strepera ) and stonechats ( Saxicola ) eat both adults and larvae. The larvae are an important food source for
4280-529: Is not divided into a pre- and postclypeus, but rather separated from the front. The antennal sclerites are fused with the surrounding head capsule, but these are sometimes separated by a suture. The number of segments in the antennae vary from six in the Accorduleceridae to 30 or more in the Pamphiliidae. The compound eyes are large with a number of facets, and there are three ocelli between
4387-401: Is two months to two years, though the adult life stage is often very short (approximately 7 – 9 days), only long enough for the females to lay their eggs. The female uses its ovipositor to drill into plant material to lay her eggs (though the family Orussoidea lay their eggs in other insects). Plant-eating sawflies most commonly are associated with leafy material but some specialize on wood, and
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4494-625: Is used to control the peach-potato aphid. Wasps RFC is an English professional rugby union team originally based in London but now playing in Coventry; the name dates from 1867 at a time when names of insects were fashionable for clubs. The club's first kit is black with yellow stripes. The club has an amateur side called Wasps FC . Among the other clubs bearing the name are a basketball club in Wantirna, Australia, and Alloa Athletic F.C. ,
4601-856: The Argidae , with some 800 species, is also worldwide, but is most common in the tropics, especially in Africa, where they feed on woody and herbaceous angiosperms . Of the other families, the Blasticotomidae and Megalodontidae are Palearctic ; the Xyelidae, Pamphilidae , Diprionidae , Cimbicidae , and Cephidae are Holarctic , while the Siricidae are mainly Holarctic with some tropical species. The parasitic Orussidae are found worldwide, mostly in tropical and subtropical regions. The wood-boring Xiphydriidae are worldwide, but most species live in
4708-623: The Royal Navy have been named HMS Wasp , the first an 8-gun sloop launched in 1749. Eleven ships of the United States Navy have similarly borne the name USS Wasp , the first a merchant schooner acquired by the Continental Navy in 1775. The eighth of these, an aircraft carrier , gained two Second World War battle stars, prompting Winston Churchill to remark "Who said a Wasp couldn't sting twice?" In
4815-620: The Triassic . The oldest superfamily, the Xyeloidea , has existed into the presents. Over 200 million years ago, a lineage of sawflies evolved a parasitoid lifestyle, with carnivorous larvae that ate the eggs or larvae of other insects. Sawflies are distributed globally, though they are more diverse in the northernmost hemispheres. The suborder name "Symphyta" derives from the Greek word symphyton , meaning 'grown together', referring to
4922-539: The comedy play Σφῆκες ( Sphēkes ), The Wasps , first put on in 422 BC. The "wasps" are the chorus of old jurors. H. G. Wells made use of giant wasps in his novel The Food of the Gods and How It Came to Earth (1904): It flew, he is convinced, within a yard of him, struck the ground, rose again, came down again perhaps thirty yards away, and rolled over with its body wriggling and its sting stabbing out and back in its last agony. He emptied both barrels into it before he ventured to go near. When he came to measure
5029-521: The evolution of eusocial behaviour . Females are diploid , meaning that they have 2n chromosomes and develop from fertilized eggs. Males, called drones, have a haploid (n) number of chromosomes and develop from an unfertilized egg. Wasps store sperm inside their body and control its release for each individual egg as it is laid; if a female wishes to produce a male egg, she simply lays the egg without fertilizing it. Therefore, under most conditions in most species, wasps have complete voluntary control over
5136-539: The head , the mesosoma (including the thorax and the first segment of the abdomen) and the metasoma. There is a narrow waist, the petiole , joining the first and second segments of the abdomen. The two pairs of membranous wings are held together by small hooks and the forewings are larger than the hind ones; in some species, the females have no wings. In females there is usually a rigid ovipositor which may be modified for injecting venom, piercing or sawing. It either extends freely or can be retracted, and may be developed into
5243-455: The masked shrew ( Sorex cinereus ), the northern short-tailed shrew ( Blarina brevicauda ) and the deer mouse ( Peromyscus maniculatus ) predate heavily on sawfly cocoons. Insects such as ants and certain species of predatory wasps ( Vespula vulgaris ) eat adult sawflies and the larvae, as do lizards and frogs. Pardalotes , honeyeaters and fantails ( Rhipidura ) occasionally consume laid eggs, and several species of beetle larvae prey on
5350-446: The 1930s and 1950s and C. pygmaeus in the 1930s and 1940s. Like all other hymenopteran insects, sawflies go through a complete metamorphosis with four distinct life stages – egg, larva, pupa and adult. Many species are parthenogenetic , meaning that females do not need fertilization to create viable eggs. Unfertilized eggs develop as male, while fertilized eggs develop into females ( arrhenotoky ). The lifespan of an individual sawfly
5457-614: The Apocrita, to form one of two suborders of Hymenoptera. Symphyta are the more primitive group, with comparatively complete venation , larvae that are largely phytophagous , and without a "wasp-waist", a symplesiomorphic feature. Together, the Symphyta make up less than 10% of hymenopteran species. While the terms sawfly and Symphyta have been used synonymously, the Symphyta have also been divided into three groups, true sawflies (phyllophaga), woodwasps or xylophaga (Siricidae), and Orussidae . The three groupings have been distinguished by
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#17327649600905564-755: The Crato Formation in Brazil, some 65 million years before the first fig trees. The Vespidae include the extinct genus Palaeovespa , seven species of which are known from the Eocene rocks of the Florissant fossil beds of Colorado and from fossilised Baltic amber in Europe. Also found in Baltic amber are crown wasps of the genus Electrostephanus . Wasps are a diverse group, estimated at well over
5671-518: The Diprionidae, such as the pine sawflies, Diprion pini and Neodiprion sertifer , cause serious damage to pines in regions such as Scandinavia. D. pini larvae defoliated 500,000 hectares (1,200,000 acres) in the largest outbreak in Finland, between 1998 and 2001. Up to 75% of the trees may die after such outbreaks, as D. pini can remove all the leaves late in the growing season, leaving
5778-870: The Second World War, a German self-propelled howitzer was named Wespe , while the British developed the Wasp flamethrower from the Bren Gun Carrier . In aerospace, the Westland Wasp was a military helicopter developed in England in 1958 and used by the Royal Navy and other navies. The AeroVironment Wasp III is a miniature UAV developed for United States Air Force special operations. Carl Eduard Adolph Gerstaecker Gerstaecker
5885-510: The Symphyta gave rise to the monophyletic suborder Apocrita ( wasps , bees , and ants ). In cladistic analyses the Orussoidea are consistently the sister group to the Apocrita. The oldest unambiguous sawfly fossils date back to the Middle or Late Triassic. These fossils, from the family Xyelidae , are the oldest of all Hymenoptera. One fossil, Archexyela ipswichensis from Queensland
5992-455: The bird family Meropidae) specialise in eating stinging insects, making aerial sallies from a perch to catch them, and removing the venom from the stinger by repeatedly brushing the prey firmly against a hard object, such as a twig. The honey buzzard attacks the nests of social hymenopterans, eating wasp larvae; it is the only known predator of the dangerous Asian giant hornet or "yak-killer" ( Vespa mandarinia ). Likewise, roadrunners are
6099-502: The body fluids of their prey. Although vespid mandibles are adapted for chewing and they appear to be feeding on the organism, they are often merely macerating it into submission. The impact of the predation of wasps on economic pests is difficult to establish. The roughly 140 species of beewolf ( Philanthinae ) hunt bees, including honeybees, to provision their nests; the adults feed on nectar and pollen. With their powerful stings and conspicuous warning coloration , social wasps are
6206-424: The chicks of several birds, including partridges . Sawfly and moth larvae form one third of the diet of nestling corn buntings ( Emberiza calandra ), with sawfly larvae being eaten more frequently on cool days. Black grouse ( Tetrao tetrix ) chicks show a strong preference for sawfly larvae. Sawfly larvae formed 43% of the diet of chestnut-backed chickadees ( Poecile rufescens ). Small carnivorous mammals such as
6313-424: The cuckoo or jewel wasps ( Chrysididae ), are kleptoparasites, laying their eggs in the nests of other wasp species to exploit their parental care. Most such species attack hosts that provide provisions for their immature stages (such as paralyzed prey items), and they either consume the provisions intended for the host larva, or wait for the host to develop and then consume it before it reaches adulthood. An example of
6420-563: The developing wasp larvae. The largest social wasp is the Asian giant hornet , at up to 5 centimetres (2.0 in) in length. The various tarantula hawk wasps are of a similar size and can overpower a spider many times its own weight, and move it to its burrow, with a sting that is excruciatingly painful to humans. The solitary giant scoliid , Megascolia procer , with a wingspan of 11.5 cm, has subspecies in Sumatra and Java ; it
6527-457: The dorsal portions of the compound eyes. The tentorium comprises the whole inner skeleton of the head. Three segments make up the thorax: the mesothorax , metathorax and prothorax , as well as the exoskeletal plates that connect with these segments. The legs have spurs on their fourth segments, the tibiae. Sawflies have two pairs of translucent wings. The fore and hind wings are locked together with hooks. Parallel development in sawfly wings
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#17327649600906634-418: The eggs have hatched, some species such as the small brown sawfly will remain with them and protects the eggs by buzzing loudly and beating her wings to deter predators. There are six larval stages that sawflies go through, lasting 2 – 4 months, but this also depends on the species. When fully grown, the larvae emerge from the trees en masse and burrow themselves into the soil to pupate. During their time outside,
6741-470: The entire suborder, there are 8,000 described species in more than 800 genera. Symphyta is paraphyletic , consisting of several basal groups within the order Hymenoptera, each one rooted inside the previous group, ending with the Apocrita which are not sawflies. The primary distinction between sawflies and the Apocrita – the ants, bees, and wasps – is that the adults lack a "wasp waist", and instead have
6848-464: The families Vespidae , Crabronidae , Sphecidae , and Pompilidae , attack and sting prey items that they use as food for their larvae; while Vespidae usually macerate their prey and feed the resulting bits directly to their brood, most predatory wasps paralyze their prey and lay eggs directly upon the bodies, and the wasp larvae consume them. Apart from collecting prey items to provision their young, many wasps are also opportunistic feeders, and will suck
6955-512: The family Vespidae contains social species, primarily in the subfamilies Vespinae and Polistinae . With their powerful stings and conspicuous warning coloration , often in black and yellow, social wasps are frequent models for Batesian mimicry by non-stinging insects, and are themselves involved in mutually beneficial Müllerian mimicry of other distasteful insects including bees and other wasps. All species of social wasps construct their nests using some form of plant fiber (mostly wood pulp) as
7062-897: The family Eulophidae attack sawflies, although their impact is low. Two species in the genus Pediobius have been studied; the two species are internal larval parasitoids and have only been found in the northern hemisphere. Parasitism of sawflies by eulophids in grass exceeds 50%, but only 5% in wheat. It is unknown as to why the attack rate in wheat is low. Furthermore, some fungal and bacterial diseases are known to infect eggs and pupa in warm wet weather. Outbreaks of certain sawfly species, such as Diprion polytomum , have led scientists to investigate and possibly collect their natural enemies to control them. Parasites of D. polytomum have been extensively investigated, showing that 31 species of hymenopterous and dipterous parasites attack it. These parasites have been used in successful biological control against pest sawflies, including Cephus cinctus throughout
7169-581: The first species to be used was Encarsia formosa , a parasitoid of a range of species of whitefly . It entered commercial use in the 1920s in Europe, was overtaken by chemical pesticides in the 1940s, and again received interest from the 1970s. Encarsia is being tested in greenhouses to control whitefly pests of tomato and cucumber , and to a lesser extent of aubergine (eggplant), flowers such as marigold , and strawberry . Several species of parasitic wasp are natural predators of aphids and can help to control them. For instance, Aphidius matricariae
7276-516: The genus Collyria . Unlike braconids, the larvae are endoparasitoids, meaning that the larvae live and feed inside the hosts body. One well known ichneumonid is Collyria coxator , which is a dominant parasitoid of C. pygmaeus . Recorded parasitism rates in Europe are between 20–76%, and as many as eight eggs can be found in a single larva, but only one Collyria individual will emerge from its host. The larva may remain inside of their host until spring, where it emerges and pupates. Several species in
7383-409: The ground. Mud daubers and pollen wasps construct mud cells in sheltered places. Potter wasps similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls. Predatory wasp species normally subdue their prey by stinging it, and then either lay their eggs on it, leaving it in place, or carry it back to their nest where an egg may be laid on
7490-693: The group's distinctive lack of a wasp waist between prostomium and peristomium. Its common name, "sawfly", derives from the saw -like ovipositor that is used for egg-laying, in which a female makes a slit in either a stem or plant leaf to deposit the eggs. The first known use of this name was in 1773. Sawflies are also known as "wood-wasps". In his original description of Hymenoptera in 1863, German zoologist Carl Gerstaecker divided them into three groups, Hymenoptera aculeata , Hymenoptera apocrita and Hymenoptera phytophaga . However, four years later in 1867, he described just two groups, H. apocrita syn. genuina and H. symphyta syn. phytophaga . Consequently,
7597-527: The historical superfamilies. Such classifications were replaced by those using molecular methods, starting with Dowton and Austin (1994). As of 2013, the Symphyta are treated as nine superfamilies (one extinct) and 25 families. Most sawflies belong to the Tenthredinoidea superfamily, with about 7,000 species worldwide. Tenthredinoidea has six families, of which Tenthredinidae is by far the largest with some 5,500 species. Extinct taxa are indicated by
7704-607: The hosts body; braconids have more of an impact on sawfly populations in the New World than they do in the Old World, possibly because there are no ichneumonid parasitoids in North America. Some braconid wasps that attack sawflies include Bracon cephi , B. lisogaster , B. terabeila and Heteropilus cephi . Female braconids locate sawfly larvae through the vibrations they produce when feeding, followed by inserting
7811-414: The larvae can be quickly dispatched by simply dropping foliage into a vessel of plain or saltwater, diluted hydrogen peroxide or isopropyl alcohol, insecticidal soap, or other garden chemical. In large-scale, industrial settings, where beneficial insect predators can also be used to eliminate larvae, as well as parasites, which have both been previously used in control programs. Small trees can be sprayed with
7918-466: The larvae exude copious amounts of salivary secretions that are avidly consumed by the adults. These include both sugars and amino acids , and may provide essential protein-building nutrients that are otherwise unavailable to the adults (who cannot digest proteins). In wasps, as in other Hymenoptera, sex is determined by a haplodiploid system, which means that females are unusually closely related to their sisters, enabling kin selection to favour
8025-403: The larvae from the leaves they are on. Larvae typically try to remain hidden on the underside of foliage. Upon removing larvae and/or the affected leaves from plants, they may be dispatched by squishing, or, alternatively, the cut leaves with larvae still attached may be fed to birds; if larger animals do not prey upon them, other insects will. However, this is not practical or useful for some, thus
8132-427: The larvae hosts; they have mouth pieces adapted to these types of feeding. Sawflies go through a complete metamorphosis with four distinct life stages – egg, larva, pupa and adult. The female uses her ovipositor to drill into plant material (or, in the case of Orussoidea, other insects) and then lays eggs in groups called rafts or pods. After hatching, larvae feed on plants, often in groups. As they approach adulthood,
8239-417: The larvae may link up to form a large colony if many other individuals are present. They gather in large groups during the day which gives them protection from potential enemies, and during the night they disperse to feed. The emergence of adults takes awhile, with some emerging anywhere between a couple months to 2 years. Some will reach the ground to form pupal chambers, but others may spin a cocoon attached to
8346-519: The larvae seek a protected spot to pupate, typically in bark or the soil. Large populations of species such as the pine sawfly can cause substantial damage to economic forestry , while others such as the iris sawfly are major pests in horticulture . Outbreaks of sawfly larvae can defoliate trees and may cause dieback, stunting or death. Sawflies can be controlled through the use of insecticides, natural predators and parasitoids, or mechanical methods. Sawflies first appeared 250 million years ago in
8453-535: The models for many species of mimic. Two common cases are Batesian mimicry , where the mimic is harmless and is essentially bluffing, and Müllerian mimicry , where the mimic is also distasteful, and the mimicry can be considered mutual. Batesian mimics of wasps include many species of hoverfly and the wasp beetle . Many species of wasp are involved in Müllerian mimicry, as are many species of bee . While wasp stings deter many potential predators, bee-eaters (in
8560-460: The name Symphyta is given to Gerstaecker as the zoological authority . In his description, Gerstaecker distinguished the two groups by the transfer of the first abdominal segment to the thorax in the Apocrita , compared to the Symphyta. Consequently, there are only eight dorsal half segments in the Apocrita, against nine in the Symphyta. The larvae are distinguished in a similar way. The Symphyta have therefore traditionally been considered, alongside
8667-474: The nineteenth-century evolution debate. The Ichneumonidae contributed to Charles Darwin 's doubts about the nature and existence of a well-meaning and all-powerful Creator. In an 1860 letter to the American naturalist Asa Gray , Darwin wrote: I own that I cannot see as plainly as others do, and as I should wish to do, evidence of design and beneficence on all sides of us. There seems to me too much misery in
8774-462: The number of prolegs and the absence of crochets in sawfly larvae. The great majority of sawflies are plant-eating , though the members of the superfamily Orussoidea are parasitic . Predators include birds, insects and small animals. The larvae of some species have anti-predator adaptations such as regurgitating irritating liquid and clustering together for safety in numbers. Sawflies are hosts to many parasitoids, most of which are Hymenoptera,
8881-432: The only real predators of tarantula hawk wasps . Social wasps are considered pests when they become excessively common, or nest close to buildings. People are most often stung in late summer and early autumn, when wasp colonies stop breeding new workers; the existing workers search for sugary foods and are more likely to come into contact with humans. Wasp nests made in or near houses, such as in roof spaces, can present
8988-408: The other female's egg from the prey and laying their own in its place. According to Emery's rule , social parasites , especially among insects, tend to parasitise species or genera to which they are closely related. For example, the social wasp Dolichovespula adulterina parasitises other members of its genus such as D. norwegica and D. arenaria . Many wasp lineages, including those in
9095-565: The outside of the host. The host remains alive until the parasitoid larvae pupate or emerge as adults. The Ichneumonidae are specialized parasitoids, often of Lepidoptera larvae deeply buried in plant tissues, which may be woody . For this purpose, they have exceptionally long ovipositors; they detect their hosts by smell and vibration. Some of the largest species, including Rhyssa persuasoria and Megarhyssa macrurus , parasitise horntails , large sawflies whose adult females also have impressively long ovipositors. Some parasitic species have
9202-507: The ovipositor can be mistaken for a stinger. Most sawflies are stubby and soft-bodied, and fly weakly. Sawflies vary in length: Urocerus gigas , which can be mistaken as a wasp due to its black-and-yellow striped body, can grow up to 20 mm ( 3 ⁄ 4 in) in length, but among the largest sawflies ever discovered was Hoplitolyda duolunica from the Mesozoic, with a body length of 55 mm ( 2 + 1 ⁄ 4 in) and
9309-581: The ovipositors of these species (such as the family Siricidae ) are specially adapted for the task of drilling through bark. Once the incision has been made, the female will lay as many as 30 to 90 eggs. Females avoid the shade when laying their eggs because the larvae develop much slower and may not even survive, and they may not also survive if they are laid on immature and glaucous leaves. Hence, female sawflies search for young adult leaves to lay their eggs on. These eggs hatch in two to eight weeks, but such duration varies by species and also by temperature. Until
9416-510: The ovipostior and paralysing the larva before laying eggs inside the host. These eggs hatch inside the larva within a few days, where they feed on the host. The entire host's body may be consumed by the braconid larvae, except for the head capsule and epidermis . The larvae complete their development within two or three weeks. Ten species of wasps in the family Ichneumonidae attack sawfly populations, although these species are usually rare. The most important parasitoids in this family are species in
9523-431: The prey item and the nest sealed, or several smaller prey items may be deposited to feed a single developing larva. Apart from providing food for their offspring, no further maternal care is given. Members of the family Chrysididae , the cuckoo wasps, are kleptoparasites and lay their eggs in the nests of unrelated host species. Like all insects, wasps have a hard exoskeleton which protects their three main body parts,
9630-569: The primary material, though this can be supplemented with mud, plant secretions (e.g., resin ), and secretions from the wasps themselves; multiple fibrous brood cells are constructed, arranged in a honeycombed pattern, and often surrounded by a larger protective envelope. Wood fibres are gathered from weathered wood, softened by chewing and mixing with saliva . The placement of nests varies from group to group; yellow jackets such as Dolichovespula media and D. sylvestris prefer to nest in trees and shrubs; Protopolybia exigua attaches its nests on
9737-626: The prolegs of both larvae gradually disappear by the time they burrow into the ground, therefore making it difficult to distinguish the two; and sawfly larvae only have a single pair of minute eyes, whereas lepidopteran larvae have four to six eyes on each side of the head. Sawfly larvae behave like lepidopteran larvae, walking about and eating foliage. Some groups have larvae that are eyeless and almost legless; these larvae make tunnels in plant tissues including wood. Many species of sawfly larvae are strikingly coloured, exhibiting colour combinations such as black and white while others are black and yellow. This
9844-673: The pupae. The larvae have several anti-predator adaptations . While adults are unable to sting, the larvae of species such as the spitfire sawfly regurgitate a distasteful irritating liquid, which makes predators such as ants avoid the larvae. In some species, the larvae cluster together, reducing their chances of being killed, and in some cases form together with their heads pointing outwards or tap their abdomens up and down. Some adults bear black and yellow markings that mimic wasps. Sawflies are hosts to many parasitoids, most of which are parasitic Hymenoptera; more than 40 species are known to attack them. However, information regarding these species
9951-418: The reproductive cycle depends on latitude ; Polistes erythrocephalus , for example, has a much longer (up to 3 months longer) cycle in temperate regions. The vast majority of wasp species are solitary insects. Having mated, the adult female forages alone and if it builds a nest, does so for the benefit of its own offspring. Some solitary wasps nest in small groups alongside others of their species, but each
10058-405: The rest being Diptera . Adult sawflies are short-lived, with a life expectancy of 7–9 days, though the larval stage can last from months to years, depending on the species. Parthenogenetic females, which do not need to mate to produce fertilised eggs, are common in the suborder, though many species have males. The adults feed on pollen, nectar, honeydew, sap, other insects, including hemolymph of
10165-432: The sex of their offspring. Experimental infection of Muscidifurax uniraptor with the bacterium Wolbachia induced thelytokous reproduction and an inability to produce fertile, viable male offspring. Females of the solitary wasp parasitoid Venturia canescens can avoid mating with their brothers through kin recognition . In experimental comparisons, the probability that a female will mate with an unrelated male
10272-602: The subfamily Polistinae . Hymenoptera in the form of Symphyta ( Xyelidae ) first appeared in the fossil record in the Lower Triassic . Apocrita, wasps in the broad sense, appeared in the Jurassic , and had diversified into many of the extant superfamilies by the Cretaceous ; they appear to have evolved from the Symphyta. Fig wasps with modern anatomical features first appeared in the Lower Cretaceous of
10379-430: The subtropical parts of Asia. Sawflies are mostly herbivores , feeding on plants that have a high concentration of chemical defences. These insects are either resistant to the chemical substances, or they avoid areas of the plant that have high concentrations of chemicals. The larvae primarily feed in groups; they are folivores , eating plants and fruits on native trees and shrubs, though some are parasitic. However, this
10486-455: The thing, he found it was twenty-seven and a half inches across its open wings, and its sting was three inches long. ... The day after, a cyclist riding, feet up, down the hill between Sevenoaks and Tonbridge, very narrowly missed running over a second of these giants that was crawling across the roadway. Wasp (1957) is a science fiction book by the English writer Eric Frank Russell ; it
10593-520: The trees too weak to survive the winter. Little damage to trees only occurs when the tree is large or when there is minimal presence of larvae. Eucalyptus trees can regenerate quickly from damage inflicted by the larvae; however, they can be substantially damaged from outbreaks, especially if they are young. The trees can be defoliated completely and may cause "dieback", stunting or even death. Sawflies are serious pests in horticulture . Different species prefer different host plants, often being specific to
10700-500: The true sawflies' ventral serrated or saw-like ovipositor for sawing holes in vegetation to deposit eggs, while the woodwasp ovipositor penetrates wood and the Orussidae behave as external parasitoids of wood-boring beetles . The woodwasps themselves are a paraphyletic ancestral grade . Despite these limitations, the terms have utility and are common in the literature. While most hymenopteran superfamilies are monophyletic , as
10807-422: The underside of leaves and branches; Polistes erythrocephalus chooses sites close to a water source. Other wasps, like Agelaia multipicta and Vespula germanica , like to nest in cavities that include holes in the ground, spaces under homes, wall cavities or in lofts. While most species of wasps have nests with multiple combs, some species, such as Apoica flavissima , only have one comb. The length of
10914-493: The wasp larvae eventually kill their hosts. Solitary wasps parasitize almost every pest insect , making wasps valuable in horticulture for biological pest control of species such as whitefly in tomatoes and other crops. Wasps first appeared in the fossil record in the Jurassic , and diversified into many surviving superfamilies by the Cretaceous . They are a successful and diverse group of insects with tens of thousands of described species; wasps have spread to all parts of
11021-402: The wasp larvae feed temporarily on their haemolymph , but if a parasitoid emerges from the host, the hyperparasites continue their life cycle inside the parasitoid. Parasitoids maintain their extreme diversity through narrow specialism. In Peru, 18 wasp species were found living on 14 fly species in only two species of Gurania climbing squash. Of the dozens of extant wasp families, only
11128-418: The world except for the polar regions. The largest social wasp is the Asian giant hornet , at up to 5 centimetres (2.0 in) in length; among the largest solitary wasps is a group of species known as tarantula hawks , along with the giant scoliid of Indonesia ( Megascolia procer ). The smallest wasps are solitary parasitoid wasps in the family Mymaridae , including the world's smallest known insect, with
11235-475: The world. I cannot persuade myself that a beneficent and omnipotent God would have designedly created the Ichneumonidae with the express intention of their feeding within the living bodies of caterpillars, or that a cat should play with mice. With its powerful sting and familiar appearance, the wasp has given its name to many ships, aircraft and military vehicles. Nine ships and one shore establishment of
11342-503: Was about twice as high as the chance of her mating with brothers. Female wasps appear to recognize siblings on the basis of a chemical signature carried or emitted by males. Sibling-mating avoidance reduces inbreeding depression that is largely due to the expression of homozygous deleterious recessive mutations. While the vast majority of wasps play no role in pollination, a few species can effectively transport pollen and pollinate several plant species. Since wasps generally do not have
11449-724: Was born in Berlin , where he studied medicine and natural sciences, receiving his PhD in 1855 as a student of Johann Christoph Friedrich Klug . In 1856 he obtained his habilitation for zoology, and soon afterwards, became a curator at the Zoological Museum of Humboldt University . In 1864 he began work as a lecturer at the Landwirtschaftlichen Lehranstalt (Agricultural Educational Facility) in Berlin. In 1874 he became an associate professor for zoology at
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