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Wasps 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. T ...
in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is
cosmopolitan Cosmopolitan may refer to: Food and drink * Cosmopolitan (cocktail), also known as a "Cosmo" History * Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953 Hotels and resorts * Cosmopoli ...
, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-nesting Ageniellini), and most capture and paralyze prey, though members of the subfamily
Ceropalinae The Ceropalinae are a subfamily of the Pompilidae Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all spe ...
are kleptoparasites of other pompilids, or ectoparasitoids of living
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s. In
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
, species may be referred to colloquially as or , though these names can be generally applied to any very large stinging wasps. Furthermore, in some parts of Venezuela and Colombia, it is called , or "horse killers", while in Brazil some particular bigger and brighter species of the general kind might be called /, or "throat locker".


Morphology

Like other strong fliers, pompilids have a
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
modified for efficient flight. The metathorax is solidly fused to the pronotum and mesothorax; moreover, the
prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum ( ventral), and the propleuron ( lateral) o ...
is best developed in Pompilidae and
Scoliidae The Scoliidae, the scoliid wasps, are a family of about 560 species found worldwide. They tend to be black, often marked with yellow or orange, and their wing tips are distinctively corrugated. Males are more slender and elongated than females, wi ...
because wasps in these families use their forelegs to dig. Pompilids typically have long, spiny legs; the hind femur is often long enough to reach past the tip of the abdomen. The tibiae of the rear legs usually have a conspicuous spine at their distal end. The first two segments of the abdomen are narrow, giving the body a slender look. The pompilid body is typically dark (black or blue, sometimes with metallic reflections), but many brightly colored species exist. From a lateral view, its pronotum looks rectangular and it extends back to the tegulae, near the base of the wings. Most species are macropterous (having long wings), but a few brachypterous (short-winged) and apterous (no wings) species are known. Spider wasps are best distinguished from other vespoid wasps in having (in most species) a transverse groove bisecting the mesopleuron (the mesepisternal sclerite, a region on the side of middle segment of the thorax above the point where the legs join). Like other Vespoidea, they have antennae with 10 flagellomeres in the female and 11 in the male. Most Pompilidae have straight inner eye margins. The hind wings do not have a distinct claval lobe, but they have a distinctive jugal lobe. The hind leg has a tibial spur with a tuft or row of fine hairs. The legs are long and slender with the tips of the tibia (metatibia) long enough to extend beyond the tip of the abdomen (metasoma).
Sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
is not pronounced, although females are often larger than the males. Coloration and wing appearance vary greatly among the many species. General coloration is
aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
(warning off predators), generally based on black, often with markings of orange, red, yellow, or white.
Larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e can also be identified by physical examination.


Systematics

The Pompilidae have in the past been split into either 4 or 6 subfamilies. However, Pitts, Wasbauer & Von Dohlen (2005) found that Notocyphinae was nested within the Pompilinae, while Epipompilinae was nested within the Ctenocerinae. This left 4 subfamilies as monophyletic clades, with Ceropalinae being the most basal subfamily and Pepsinae being the sister clade to the more derived Ctenocerinae and Pompilinae. Waichert et al (2015) resurrected the Notocyphinae as a subfamily, to have 5 subfamilies within Pompilidae. In this classification Ctenocerinae were found to be the basal group; Pepsinae and Notocyphinae were sister taxa, as were Pepsinae and Pompilinae. They placed the genus ''
Epipompilus ''Epipompilus'' is a genus of spider wasps in the subfamily Pepsinae, part of the widespread family Pompilidae. Representatives of ''Epipompilus'' can be found in Australasia and North and South America. This distribution may indicate that ''Ep ...
'' in the Pepsinae. The subfamilies according to Waichert et al are: *
Ceropalinae The Ceropalinae are a subfamily of the Pompilidae Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all spe ...
2 genera, ''
Ceropales ''Ceropales'' is a genus of kleptoparasitic spider wasps from the sub-family Ceropalinae of the family Pompilidae. They are characterised by the taking of the spider prey of other solitary wasps, mainly Pompilidae but members of the Sphecidae ...
'' and ''
Irenangelus ''Ireangelus'' is a genus of kleptoparasitic spider wasps from the sub-family Ceropalinae of the family Pompilidae. The genus has a pan tropical distribution, being known from Oriental, Neotropical, Australian, eastern Palearctic, and Madagas ...
'' * Notocyphinae containing a single genus, ''
Notocyphus ''Notocyphus'' is a genus of spider wasps, belonging to the family Pompilidae. They are the only genus in the monotypic subfamily Notocyphinae. These wasps are found in the Nearctic and the Neotropics. Species The following species are included ...
'', distributed in both the
Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...
and
Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
*
Ctenocerinae The Ctenocerinae are a subfamily of spider wasps, Pompilidae, which contains a small number of genera, two in the Neotropics, four in Australia and the remainder in Africa. Ctenocerine wasps have evidently evolved from a common ancestor with the ...
containing 2 genera in the Neotropics, 4 in Australia and 11 in Africa *
Pepsinae The Pepsinae are a subfamily of the spider wasp family, Pompilidae, including the tarantula hawks, as well as smaller species. Genera *'' Ageniella'' Banks, 1912 *'' Allaporus'' Banks, 1933 *'' Auplopus'' Spinola, 1841 250px, '' Auplopus carbo ...
a diverse subfamily distributed worldwide except for Antarctica *
Pompilinae The Pompilinae are a subfamily of the spider wasp family, Pompilidae Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfam ...
a diverse subfamily distributed worldwide except for Antarctica


Evolution

The oldest fossil currently known is an indeterminate fossil from the Early
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
(
Ypresian In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian ...
)
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Wash ...
of Washington State, USA. Other fossil species are known from Dominican and Baltic ambers, the
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones an ...
and various other localities in Germany, France and Spain. '' Bryopompilus'' described from the mid Cretaceous
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
was initially thought to belong to this family, however it was subsequently placed in its own family, the Bryopompilidae.


Ecology and behavior

Unlike many other families in the
Aculeata Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot ...
, essentially all wasps in this family are solitary (nests made by a single female). Adult pompilids are nectar-feeding insects and feed on a variety of plants. Depending on genus and species, pompilids capture a variety of spiders for their larvae to feed on, covering nearly all free-living spider families, including tarantulas,
wolf spider Wolf spiders are members of the family Lycosidae (). They are robust and agile hunters with excellent eyesight. They live mostly in solitude, hunt alone, and do not spin webs. Some are opportunistic hunters, pouncing upon prey as they find it or ...
s (Lycosidae), huntsman spiders (Sparassidae),
jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
s (Salticidae) and baboon spiders (Harpactirinae), though any given pompilid tends to attack only a limited diversity of spiders. A female wasp searches the ground and/or vegetation for a spider, and upon finding one, stings it, paralyzing the spider. The targeted spider is typically unable to kill the wasp, because the wasp can just fly out of reach, so at best the spider fights fiercely to escape. Tarantula hawks (Pepsini) do not attack when adult tarantulas are close to or in their burrows. Instead, the wasps seek out adult males who have left their burrows in search of females to mate with. In the open the wasp first uses its wings to beat air over the tarantula, deceiving the tarantula into thinking that it is being targeted by a large bird so the tarantula reacts by curling up to appear smaller and less noticeable, which in turn makes the tarantula defenseless against the wasp's attack. However, Brazilian Wandering spiders (''
Phoneutria ''Phoneutria'' is a genus of spiders in the family Ctenidae. Phoneutria venom is potentially medically significant to humans. They are mainly found in northern South America, with one species in Central America. Members of the genus are commo ...
'') and their predators have a different interaction dynamic, and the spiders often manage to defeat the hunting wasp. Once the spider is paralyzed, a female pompilid makes a burrow or flies or drags the spider to a previously made burrow.Spider Wasps
Australian Museum Online
Because of the large body size of their prey, tarantula hawks usually will either construct burrows near the site of attack or use the host's own burrow or tunnel. Pompilids typically provide each of their larvae with a single prey/host, which must be large enough to serve as its food source throughout its development. Typically, a single egg is laid on the abdomen of the spider, and the nest or burrow is closed so the larva can develop without disruption by other parasites or scavengers. The female wasp may then engage in spreading soil or other changes to the area, leaving the nest site inconspicuous. One species of spider wasp protects its nests by putting dead ants into the outermost chamber, where the ants' chemicals deter predators. The egg hatches and the larva feeds on the spider, breaking through the integument with its mandibles. As the larva feeds on its host, it saves the vital organs, such as the heart and central nervous system, for last. By waiting until the final larval instar, it ensures the spider will not decompose before the larva has fully developed. The larva has five
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
stages before it pupates; no major morphological differences are noted between the first four instars, with the exception of size. At the conclusion of the final instar, the larva spins a durable silk cocoon, and emerges as an adult either later in the same season or overwinters, depending on the species and the time of year the larva pupates. Some ceropalines lay their egg on a still-active spider, only temporarily paralyzing it, and the wasp larva feeds externally by extracting
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
after the egg hatches. In time, the spider will die, and the mature wasp larva will then pupate. The size of the host can influence whether the wasp's egg will develop as a male or a female; larger prey often yield the (larger) females. ''Pepsis thisbe'' of the southwestern United States exhibits a direct correlation between adult wasp body length and the weight of its host spider, ''Aphonopelma echina''. Because the size of a ''P. thisbe'' adult is determined by the size of the host provided for it by its mother, the seasonal frequency of host sizes implicitly will determine the size variation in adult wasps. In another study on ''Pepsis thisbe'', chemosensory cues were shown to be used to detect specific hosts. Specific chemosensory cues attract the wasp to its prey, ''Aphonopelma echina'', despite other host spiders of the same size and frequency being present. In studies on ''Pepsis grossa'' (formerly ''P. formosa''), a pompilid of the southwestern United States, the wasps were found to have behavioral plasticity. Their hunting behavior concerning their host ''Rhechostica echina'' improved with experience. The time required to complete all behavioral components decreased with each spider killed. Concerning mating behavior, males acquire perch territories to scan for incoming receptive females. In studies on the tarantula wasp ''Hemipepsis ustulata'', larger males are more likely to acquire perch territories and territorial males appear to increase their chances of mating because receptive females fly to perch sites held by said males.


Sting


Toxins

The Pompilidae produce a venom, delivered when they sting, containing a variety of powerful
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature nerv ...
s named pompilidotoxin (PMTX). These inhibit the inactivation of voltage-gated sodium channels, causing too much sodium to flow through neuron cell membranes, causing long bursts of
nerve impulse An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, c ...
s (action potentials), and thus overstimulating these nerves. Alpha- and beta-PMTX are both small
peptide Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. ...
chains of just 13
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s; alpha-PMTX has been studied in '' Anoplius samariensis'', while beta-PMTX has been studied in '' Batozonellus maculifrons''.


Schmidt pain index

In 1984, Justin O. Schmidt developed a hymenopteran sting pain scale, now known as the
Schmidt sting pain index The Schmidt sting pain index is a pain scale rating the relative pain caused by different hymenopteran stings. It is mainly the work of Justin O. Schmidt (born 1947), an entomologist at the Carl Hayden Bee Research Center in Arizona, United Sta ...
. In this index, a 0 is given to a sting from an insect that cannot break through human skin, a 2 is given for intermediate pain, and a 4 is given for intense pain. The scale rates stings from 78 different species in 42 different genera. Spider wasps of the genus ''Pepsis'', also known as tarantula hawks, have a sting rating of 4. The sting is described as "blinding, fierce, and shockingly electric. A running hair dryer has been dropped into your bubble bath." Only the sting of the bullet ant, ''Paraponera clavata'', is ranked higher, with a 4+ rating.


Gallery

Image:Spider wasp flying.jpg, Spider wasp flying with prey Image:Anoplius viaticus.jpg, ''Anoplius viaticus'' (L.) (Hymenoptera: Pompilidae) with prey ''Nuctenea umbratica'' (Clerck) (Arachnida: Araneidae), England UK Image:Wasp-Spider Hunting.jpg, '' Tachypompilus ferrugineus'', also known as the rusty spider wasp, hunting in
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
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Image:Spider wasp 05.JPG, Spider wasp with its prey near
Heemstede Heemstede () is a town and a municipality in the Netherlands, in the province of North Holland. It is the fourth richest municipality of the Netherlands. History Heemstede formed around the Castle ''Heemstede'' that was built overlooking the ...
,
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
Image:Spider Wasp 003.jpg, Spider wasp attacking huntsman spider in
Sydney Sydney ( ) is the capital city of the state of New South Wales, and the most populous city in both Australia and Oceania. Located on Australia's east coast, the metropolis surrounds Sydney Harbour and extends about towards the Blue Mounta ...
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Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
Image:Wasp Ants.jpg, Spider wasp being carried off by ants. Image:Spider-wasp.jpg, A pompilid from
Bangalore Bangalore (), officially Bengaluru (), is the capital and largest city of the Indian state of Karnataka. It has a population of more than and a metropolitan population of around , making it the third most populous city and fifth most ...
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Image:Spiderwasp1 feb09.jpg, A Western Australian pompilid has captured a large huntsman spider. Image:Pompilidae en Quito Ecuador.jpg, A pompilid has captured a spider in
Quito Quito (; qu, Kitu), formally San Francisco de Quito, is the capital and largest city of Ecuador, with an estimated population of 2.8 million in its urban area. It is also the capital of the province of Pichincha. Quito is located in a valley on ...
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. Image:Spiderwasp2 feb09.jpg, The anaesthetised spider is carried up to a nest in the roof. Image:Spider wasp pompilid.jpg, A spider wasp is dragging its captured prey in
Sydney Sydney ( ) is the capital city of the state of New South Wales, and the most populous city in both Australia and Oceania. Located on Australia's east coast, the metropolis surrounds Sydney Harbour and extends about towards the Blue Mounta ...
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Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
. Image:IndianSpiderWasp.JPG, An Indian spider wasp carrying a jumping spider in
Tumkur Tumkur, officially renamed as Tumakuru, is a city located in the southern part of Indian state of Karnataka. Tumkur is situated at a distance of northwest of Bangalore, the state capital along NH 48 and NH 73. It is the headquarters of the ...
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Image:Spider Wasp In Nairobi.jpg, A spider wasp feeding on a spider in
Nairobi Nairobi ( ) is the capital and largest city of Kenya. The name is derived from the Maasai phrase ''Enkare Nairobi'', which translates to "place of cool waters", a reference to the Nairobi River which flows through the city. The city proper ...
, Kenya.


References


External links

* * * A series of photos showing ''Anoplius viaticus'' trapping a spider and fighting off other predators, in order to lay her egg. * Australian government agency fact sheet. * {{Taxonbar, from=Q252859 Articles containing video clips