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| {{Short description|Vespid wasps that gather fibers from dead wood and plant stems}} | |||
| Despite the use of the proper name, most social ]s make nests from paper, although some tropical wasp species such as ''Listenogaster Flavolineata'' use mud, a far more easy resource for the wasp to collect. The larger colonial species, ]s, ]s, can be very defensive and should not be approached unless one is experienced. All are beneficial and should be protected if possible. In ], hornets are a threatened species and are protected by law. | |||
| {{Paraphyletic group | |||
| | auto = yes | |||
| | image = Wasp colony.jpg | |||
| | image_caption = A young paper wasp queen ('']'') starting a new colony | |||
| | parent = Vespidae | |||
| | includes_text = Subfamilies with paper wasp species | |||
| | includes = *] | |||
| *] | |||
| *] | |||
| }} | |||
| ]); exposed comb]] | |||
| ] | |||
| ] nest (]); concealed comb]] | |||
| '''Paper wasps''' are a type of ] ]s. The term is typically used to refer to members of the vespid subfamily ], though it often colloquially includes members of the subfamilies ] (]s and ]s) and ], which also make nests out of paper. | |||
| Paper wasps gather fibers from old wood and dead dry plant stems, chew them and mix with ] to make a water-resistant gray paper. The nests are characterised by having open cells and a ] attaching the nest to the substrate. The wasps produce a chemical which repels ants and secretes it around the base of this petiole in order to avoid letting their brood become ant food. Humans may have learned to make paper from watching wasps. In the photo one can see a much roughed deck post from the continual gathering of wood fibers by wasps making nests. | |||
| Paper wasp nests are characterized by open combs with down pointing cells. Some types of paper wasps are occasionally referred to as '''umbrella wasps''' due to the distinctive design of their nests.<ref name=enc> Microsoft Encarta Online Encyclopedia 2006.</ref> | |||
| The European Polistes wasp, ''Polistes dominulus'' (the species in the upper photo) was introduced into the ] about ] and has quickly spread throughout most of the country, in most cases replacing native species within a couple years. It nests earlier in the spring, in a wider variety of nest sites, is more aggressive, and feeds on a larger variety of insects. Native species feed almost entirely on caterpillars. Some entomologists are considering this to be an ], while others note that it may become an important control of ], a serious invasive pest. | |||
| == Species == | |||
| Approximately 300 species of '']'' paper wasps have been identified worldwide. The most common paper wasp in Europe is '']''.<ref>{{cite web |url=http://www.ipc.ibaraki.ac.jp/~jkrte/wasp/polistes/Polistes.htm |title=Distributional checklist of the species of the genus ''Polistes'' (Hymenoptera: Vespidae; Polistinae, Polistini) |author=James M. Carpenter |access-date=2009-02-09 |url-status=dead |archive-url=https://web.archive.org/web/20090710131558/http://www.ipc.ibaraki.ac.jp/~jkrte/wasp/polistes/Polistes.htm |archive-date=2009-07-10 }}</ref> The ] tribe ] contains another 300 species, and the ] tribes ] and ] each contain over 250 more, so the total number of true paper wasps worldwide is about 1100 species, almost half of which can be found in the neotropics. | |||
| ] fibers from a ] in ]]] | |||
| The general life cycle of Polistes follows a cycle that can be divided into four phases (after Reeve 1991): | |||
| == Nests == | |||
| #Founding (or pre-emergence) phase | |||
| ]'' on part of a large nest, in California]] | |||
| #Worker phase | |||
| #Reproductive phase | |||
| #“Intermediate” phase | |||
| The nests of most true paper wasps are characterized by having open combs with cells for brood rearing. The wasps gather ] from dead ] and ], which they mix with ], and use to construct nests that appear made out of gray or brown papery material. A ']', or constricted stalk attaches the nest to a branch or other structure.<ref name=OSU>Lyon, W.F. and G.S. Wegner (1991). {{webarchive|url=https://web.archive.org/web/20010425015741/http://ohioline.osu.edu/hyg-fact/2000/2077.html |date=2001-04-25 }} Ohio State University Extension Fact Sheet: Entomology</ref> | |||
| The founding stage involves a solitary female initiating a nest, building 20-30 cells before initially ovipositing. This phase begins in spring, depending on climatic conditions. The foundress or foundresses begin by fashioning a ] and produce a single cell at the end of it. Further cells are then added around this, 6 cells surrounding it to produce the characteristic hexagonal shape of the cells. | |||
| The wasps secrete an ] repellent chemical which they spread around the base of the petiole or anchor to prevent the loss of eggs or ].<ref>{{cite journal |last1=Post |first1=David C |last2=Jeanne |first2=Robert L |title=Colony Defense against Ants by Polistes fuscatus (Hymenoptera: Vespidae) in Wisconsin |journal=Journal of the Kansas Entomological Society |date=July 1981 |volume=54 |issue=3 |page=599 |jstor=25084198 |url=https://www.jstor.org/stable/25084198 |access-date=18 December 2021}}</ref> | |||
| ] | |||
| Most social ]s of the family Vespidae make nests from paper, but some ] species, such as '']'', use ]. A small group of ] ] wasps, of the genus '']'' (the only eusocial wasps outside the family Vespidae), also constructs nests out of chewed plant fibers, though the nest consistency is quite different from those of true paper wasps, due to the absence of wood fibers, and the use of silk extruded by female wasps to bind the fibers.<ref>{{cite journal |first1=R. W. |last1=Matthews |first2=C. K. |last2=Starr |title=Microstigmus comes Wasps have a Method of Nest Construction Unique Among Social Insects |journal=Biotropica |volume=16 |issue=1 |year=1984 |pages=55–58 |doi=10.2307/2387895 |jstor=2387895 |bibcode=1984Biotr..16...55M }}</ref> | |||
| ] | |||
| Nests can be found in sheltered areas, such as the eaves of a house, the branches of a tree, on the end of an open pipe, or on an old clothesline. Some species, such as '']'', will vary their nest architecture depending on where they build their nest.<ref name=":2">{{Cite journal|title = Nest Architecture of the Australian Paper Wasp Ropalidia Romandi Cabeti, With a Note on its Developmental Process (Hymenoptera: Vespidae)|last1 = Yamane|first1 = Soichi|date = 1994|journal = Psyche: A Journal of Entomology|doi =10.1155/1994/92839 |last2 = Ito|first2 = Yosiaki|issue = 3–4|pages = 145–158|volume = 101|doi-access = free}}</ref> | |||
| After the hatching of the first larvae, the foundress progressively provisions (brings food multiple times throughout development) them with malaxated caterpillar flesh, halting further oviposition until some of the larvae have pupated. These larvae will eventually hatch to become 1st generation ]. | |||
| Three species of '']'' are obligate ], and have lost the ability to build their own nests, and are sometimes referred to as "]s". They rely on the nests of their hosts to raise their brood.<ref name="Dapporto">Dapporto L, Cervo R, Sledge MF, Turillazzi S (2004) "Rank integration in dominance hierarchies of host colonies by the paper wasp social parasite ''Polistes sulcifer'' (Hymenoptera, Vespidae)". ''J Insect Physiol'' '''50''' :217–223</ref> A few ]s and ]s are also brood parasites (e.g., '']'').<ref>{{Cite web|url = http://www.bwars.com/index.php?q=wasp/vespidae/vespinae/vespula-austriaca|title = Vespula austriaca (Panzer, 1799)|date = 1998|access-date = 15 Oct 2014|website = Bees, Wasps & Ants Recording Society|publisher = BWARS|last = Archer|first = M.E.}}</ref> | |||
| As well as founding a nest, a female can also choose to attempt to usurp a nest of a ], or join conspecifics upon another nest. In the case of the latter, evidence shows that such co-founding females are generally, but not exclusively, closely related (reviewed in Reeve 1991). | |||
| == Brood rearing == | |||
| The worker phase usually begins in early summer, roughly 2 months after colony initiation, with the emergence of the first workers. On emergence, the workers take up most of the colony’s foraging, brood care and nest maintenance. Typically at this stage, the auxillary (subordinate) foundresses are driven from the nest, leaving the alpha female and newly emerged workers. | |||
| In temperate climates, in the fall, reproductive males and females are reared. They fly away and mate. Only the new queens hibernate overwinter in a sheltered location. | |||
| In the spring each searches for a suitable location and starts and maintains a new nest typically by herself until the first brood of female workers matures and helps in all activities except egg laying. | |||
| ] | |||
| ] | |||
| ] | |||
| Once eggs hatch, the legless larvae remain in their cells to be taken care of. The wasps hunt prey, primarily ]s, but also other insects like ], and ] ]. The adult cuts up the prey and masticates it (thoroughly chewing it) prior to feeding it to the larvae. | |||
| Emergence of the first female reproductives (Gynes) is taken to signal the start of the reproductive phase of the colony. The reproductives differ from workers produced at this latter phase of the colony by having increased levels of fats stores to allow it to survive the over-wintering period, as well as having increased levels of cryoprotectant carbohydrate compounds for the same purpose. | |||
| ] | |||
| Water is also provided for the larva. Wasps may find a puddle or stream or dew that has collected at the base of some leaves. | |||
| Finally, the “intermediate” phase is really just the period in which the gynes and males mate and then disperse from the natal colony, before over wintering (hibernating) until the start of the next colony cycle. Typically this period is characterised by a decline in brood care and foraging, as well as fewer workers (due to mortality without the worker being replaced by newly hatched brood). In P. dominulus the colony disperses in the late summer. | |||
| ] | |||
| When the larvae mature, they pupate in their cell and the workers cap the cell, sealing them in until the adult breaks out of its pupal shell. | |||
| ===Dominance hierarchy system=== | |||
| The open nest is light colored reflecting light and the nest material provides some insulation, even so it can be subjected to elevated temperatures with sun exposure or high ambient temperatures. The wasps can cool their nest by fanning their wings and/or bringing water to place in the nest letting evaporation provide substantial cooling. | |||
| Morphologically, there is little difference between the foundress and subordinate reproductive members of the colony. However, several studies have shown that behavioural differentiation occurs (Theraulaz 1992, Pardi 1942), the role the individual female taking determined by social interaction within the colony. Typically, the alpha female dominates all other individuals of a colony, and this female lays the majority of eggs, and partakes in differential oophagy. The alpha female devotes much of her time to social interaction, in comparison to subordinates that are much more involved in foraging and brood care (Theraulaz 1989). | |||
| ] | |||
| ] | |||
| == Behavior == | |||
| These behavioural divisions are not permanent; if an alpha female is removed from a nest then another female (usually the second-most dominant, beta female) assumes the role and behavioural profile of the removed dominant. Indeed, individuals alternate between different profiles of behaviour within their own dominance rank position. | |||
| Unlike ]s and ]s, which can be very aggressive, ] paper wasps will generally only attack if they themselves or their nest are threatened.<ref>Felixson, Carol (undated). "Paper wasps work together." Retrieved 2009-04-26 from "L.A. Times" at .</ref> Their territoriality can lead to attacks on people, and their stings are quite painful and – like all venomous animals – can produce a potentially fatal ] reaction in some individuals.<ref name=Texas>Drees, B.M. and John Jackman (1999). ''Field Guide to Texas Insects.'' Gulf Publishing Company, Houston, Texas. Excerpt available at: {{webarchive|url=https://web.archive.org/web/20061205022343/http://insects.tamu.edu/fieldguide/cimg348.html |date=2006-12-05 }}</ref> A study conducted on European paper wasps ('']'') concluded that wasps with brighter ] are more venomous, because they have larger venom glands, and offer a stronger warning signal to organisms threatening the nest.<ref>{{Cite journal|last1=Vidal-Cordero|first1=J|last2=Moreno-Rueda|first2=Gregorio|last3=López-Orta|first3=Antonio|last4=Marfil-Daza|first4=Carlos|last5=Ros-Santaella|first5=José L|last6=Ortiz-Sánchez|first6=F|date=2012|title=Brighter-colored paper wasps (''Polistes dominula'') have larger poison glands |journal=Frontiers in Zoology | volume=9|issue=1|pages=20|doi=10.1186/1742-9994-9-20|issn=1742-9994|pmc=3495029|pmid=22901602|doi-access=free}}</ref> | |||
| ] | |||
| Some studies (Pardi 1946) seem to indicate that the dominant female, through its behaviour, suppresses the ovarian development of subordinates. Abdominal wagging is thought to serve as a dominance signal between dominant foundresses and subordinates, but studies by Roseler and Roseler (1989) showed that ovariectomiszed dominants failed to restrict subordinate reproduction whilst still retaining dominance. | |||
| ] | |||
| ] | |||
| Most wasps are beneficial in their natural habitat and are critically important in natural biocontrol.<ref name= OSU/> Paper wasps feed on sugars like ], ] ] and the sugary liquid produced by their larvae. Because they are a known ] and feed on known garden ], paper wasps are often considered to be beneficial by gardeners.<ref name=Texas/> | |||
| ===Nestmate recognition=== | |||
| ] of paper wasps huddling together in early winter.]] | |||
| ''Polistes'' discriminate colony mates using an acquired (i.e. learned) cue, absorbing hydrocarbons from the natal nest at eclosion (Gamboa 1996). This cuticular hydrocarbon "signature" is derived both from the plant material and the foundress-applied substances that the nest is made of. Studies of ''Polistes fucatus'' have looked into the molecular basis of the recognition "pheromone" used by the wasps, and indicate that at least some of the recognisable labels have the same chemical constituents as the adult cuticular hydrocarbons. | |||
| Dominant individuals of ''P. dominulus'' have differing cuticular profile to workers (Bonavita-Cougourdan 1991), and the frequent observations of the dominant female stroking its gasters across the nest surface, combined with its staying on the nest for longer times than subordinates, suggests that the dominant individual may contribute more to the nest odour. | |||
| When threatened, the wasps have a variety of responses depending upon the severity. Paper wasps use ]s to coordinate their response. The first level is posturing. They face the perceived threat, stand tall and raise their wings. | |||
| A study of P. Carolina showed that females do not preferentially feed their own progeny (Strassman 2000) (as larvae), so it may be the case that nest odour only serves as a likely indicator of relatedness, rather than a specific label of kinship. | |||
| At a higher level of threat, the wasp will move around the nest surface. With sufficient disturbance, the wasp will fly around the nest attempting to locate the source, chasing and stinging the threat.{{cn|date=September 2024}} | |||
| Further to this recognition of nest-mates, a study on Polistes biglumis illustrated how foundresses discriminate between ‘alien’ eggs and their own, via differential oophagy (Lorenzi 2000). Interestingly, the discrimination focused upon eggs destined to be reproductives, with ‘alien’ worker destined eggs allowed to remain on the nest. The authors speculated that the benefits of allowing worker destined eggs to remain (and so hatch to become workers which will then aid the colony) outweigh the costs of initially provisioning the resultant larvae. | |||
| == See also == | |||
| The mechanism of differentiation was not elucidated, but was thought to be based upon differences in cuticular hydrocarbon odour. Whether the discriminatory oophagy was a result of decreased tolerance of alien odours during the later, reproductive phase of the colony cycle, or an actual discrimination between worker and reproductive destined eggs, remains to be supported with good evidence. | |||
| * ] | |||
| * ] | |||
| * '']'' | |||
| * ] | |||
| * ] | |||
| == |
== References == | ||
| {{Reflist}} | |||
| #Nest building in a Social Wasp: Postures and Constraints (Hymenoptera: Vespidae). Karsai I. & Theraulaz G., Sociology Vol. 26, No.1 pp. 83-114, 1995. | |||
| #Intra-specific variation in the comb structure of Polistes dominiculus: parameters, maturation, nest size and cell arrangement. Karsai I. & Penzes Z., Insect Sociaux Vol. 43 pp. 277-296, 1996. | |||
| #''Polistes''. Reeve H. K., in "The Social Biology of Wasps", edited by Ross K. G. & Mathews R. W., pp. 99-148, 1991. | |||
| #The dynamics of colony organisation in the primitively eusocial wasp Polistes dominulus Christ. Theraulaz G., Gervet J. et al. Ethology 91, pp. 177-202, 1992. | |||
| #Pardi (1942, 1946) from table compiled in Ito Y.: "Behaviour and Social Evolution of Wasps: The Communal Aggregation Hypothesis", p. 46 1993. | |||
| #Effects of removal of alpha individuals from a Polistes dominulus Christ. Wasp society: Changes in behavioural patterns resulting from hierarchical changes. Theraulaz G., Pratte M. & Gervet J.; Insectes Sociaux 5 pp. 169-179 1989. | |||
| #Kin recognition pheromones in social wasps: combining chemical and behavioural evidence. Gamboa G. J., Grudzien T.A., Espelie K.E. & Bura E.A. Animal Behaviour 51, pp.625-629 1996. | |||
| #Cuticular hydrocarbons, social organisation and ovarian development in a polistine wasp: Polistes dominulus. Bonavita-Cougourdan A., Theraulaz G., Bagneres A.G., Roux M., Pratte M., Provost E., Clement J.L.; Comp. Biochem. Physiol. B Biochem. Mol. Biol. 100 pp. 667-680 1991. | |||
| #Absence of within-colony kin discrimination: foundresses of the social wasp, Polistes Carolina, do not prefer their own larvae. Strassman J. E., Seppa P. & Queller D.C.; Naturwissenschaften 87 pp.266-269 2000. | |||
| #Opportunistic discrimination of alien eggs by social wasps (Polistes biglumis, Hymenoptera Vespidae): a defence against social parasitism? Lorenzi M. C. & Filippone F.; Behav. Ecol. Sociobiol 48: pp.402-406 2000. | |||
| == See also == | |||
| * ] | |||
| * '']'' | |||
| == External links == | |||
| ] | |||
| * Retrieved 2012-12-04. | |||
| ] | |||
| ] | |||
| ] | |||
| ] | |||
| ] | |||
| ] | |||
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Latest revision as of 20:06, 10 October 2024
Vespid wasps that gather fibers from dead wood and plant stems
| Paper wasp | |
|---|---|
| |
| A young paper wasp queen (Polistes gallicus) starting a new colony | |
Scientific classification
| |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Hymenoptera |
| Superfamily: | Vespoidea |
| Family: | Vespidae |
| Subfamilies with paper wasp species | |
Paper wasps are a type of vespid wasps. The term is typically used to refer to members of the vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae (hornets and yellowjackets) and Stenogastrinae, which also make nests out of paper.
Paper wasp nests are characterized by open combs with down pointing cells. Some types of paper wasps are occasionally referred to as umbrella wasps due to the distinctive design of their nests.
Species
Approximately 300 species of Polistes paper wasps have been identified worldwide. The most common paper wasp in Europe is Polistes dominula. The Old World tribe Ropalidiini contains another 300 species, and the Neotropical tribes Epiponini and Mischocyttarini each contain over 250 more, so the total number of true paper wasps worldwide is about 1100 species, almost half of which can be found in the neotropics.
Nests
The nests of most true paper wasps are characterized by having open combs with cells for brood rearing. The wasps gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests that appear made out of gray or brown papery material. A 'petiole', or constricted stalk attaches the nest to a branch or other structure. The wasps secrete an ant repellent chemical which they spread around the base of the petiole or anchor to prevent the loss of eggs or brood.
Most social wasps of the family Vespidae make nests from paper, but some stenogastrine species, such as Liostenogaster flavolineata, use mud. A small group of eusocial crabronid wasps, of the genus Microstigmus (the only eusocial wasps outside the family Vespidae), also constructs nests out of chewed plant fibers, though the nest consistency is quite different from those of true paper wasps, due to the absence of wood fibers, and the use of silk extruded by female wasps to bind the fibers.
Nests can be found in sheltered areas, such as the eaves of a house, the branches of a tree, on the end of an open pipe, or on an old clothesline. Some species, such as Ropalidia romandi, will vary their nest architecture depending on where they build their nest.
Three species of Polistes are obligate social parasites, and have lost the ability to build their own nests, and are sometimes referred to as "cuckoo paper wasps". They rely on the nests of their hosts to raise their brood. A few hornets and yellowjackets are also brood parasites (e.g., Vespula austriaca).
Brood rearing
In temperate climates, in the fall, reproductive males and females are reared. They fly away and mate. Only the new queens hibernate overwinter in a sheltered location. In the spring each searches for a suitable location and starts and maintains a new nest typically by herself until the first brood of female workers matures and helps in all activities except egg laying.
Once eggs hatch, the legless larvae remain in their cells to be taken care of. The wasps hunt prey, primarily caterpillars, but also other insects like flies, and beetle larvae. The adult cuts up the prey and masticates it (thoroughly chewing it) prior to feeding it to the larvae.
Water is also provided for the larva. Wasps may find a puddle or stream or dew that has collected at the base of some leaves.
When the larvae mature, they pupate in their cell and the workers cap the cell, sealing them in until the adult breaks out of its pupal shell. The open nest is light colored reflecting light and the nest material provides some insulation, even so it can be subjected to elevated temperatures with sun exposure or high ambient temperatures. The wasps can cool their nest by fanning their wings and/or bringing water to place in the nest letting evaporation provide substantial cooling.
Behavior
Unlike yellowjackets and hornets, which can be very aggressive, polistine paper wasps will generally only attack if they themselves or their nest are threatened. Their territoriality can lead to attacks on people, and their stings are quite painful and – like all venomous animals – can produce a potentially fatal anaphylactic reaction in some individuals. A study conducted on European paper wasps (Polistes dominula) concluded that wasps with brighter aposematic colors are more venomous, because they have larger venom glands, and offer a stronger warning signal to organisms threatening the nest.
Most wasps are beneficial in their natural habitat and are critically important in natural biocontrol. Paper wasps feed on sugars like nectar, aphid honeydew and the sugary liquid produced by their larvae. Because they are a known pollinator and feed on known garden pests, paper wasps are often considered to be beneficial by gardeners.
When threatened, the wasps have a variety of responses depending upon the severity. Paper wasps use alarm pheromones to coordinate their response. The first level is posturing. They face the perceived threat, stand tall and raise their wings.
At a higher level of threat, the wasp will move around the nest surface. With sufficient disturbance, the wasp will fly around the nest attempting to locate the source, chasing and stinging the threat.
See also
References
- "Paper Wasp" Microsoft Encarta Online Encyclopedia 2006.
- James M. Carpenter. "Distributional checklist of the species of the genus Polistes (Hymenoptera: Vespidae; Polistinae, Polistini)". Archived from the original on 2009-07-10. Retrieved 2009-02-09.
- ^ Lyon, W.F. and G.S. Wegner (1991). Paper Wasps and Hornets Archived 2001-04-25 at the Wayback Machine Ohio State University Extension Fact Sheet: Entomology
- Post, David C; Jeanne, Robert L (July 1981). "Colony Defense against Ants by Polistes fuscatus (Hymenoptera: Vespidae) in Wisconsin". Journal of the Kansas Entomological Society. 54 (3): 599. JSTOR 25084198. Retrieved 18 December 2021.
- Matthews, R. W.; Starr, C. K. (1984). "Microstigmus comes Wasps have a Method of Nest Construction Unique Among Social Insects". Biotropica. 16 (1): 55–58. Bibcode:1984Biotr..16...55M. doi:10.2307/2387895. JSTOR 2387895.
- Yamane, Soichi; Ito, Yosiaki (1994). "Nest Architecture of the Australian Paper Wasp Ropalidia Romandi Cabeti, With a Note on its Developmental Process (Hymenoptera: Vespidae)". Psyche: A Journal of Entomology. 101 (3–4): 145–158. doi:10.1155/1994/92839.
- Dapporto L, Cervo R, Sledge MF, Turillazzi S (2004) "Rank integration in dominance hierarchies of host colonies by the paper wasp social parasite Polistes sulcifer (Hymenoptera, Vespidae)". J Insect Physiol 50 :217–223
- Archer, M.E. (1998). "Vespula austriaca (Panzer, 1799)". Bees, Wasps & Ants Recording Society. BWARS. Retrieved 15 Oct 2014.
- Felixson, Carol (undated). "Paper wasps work together." Retrieved 2009-04-26 from "L.A. Times" at .
- ^ Drees, B.M. and John Jackman (1999). Field Guide to Texas Insects. Gulf Publishing Company, Houston, Texas. Excerpt available at: Texas Cooperative extension Archived 2006-12-05 at the Wayback Machine
- Vidal-Cordero, J; Moreno-Rueda, Gregorio; López-Orta, Antonio; Marfil-Daza, Carlos; Ros-Santaella, José L; Ortiz-Sánchez, F (2012). "Brighter-colored paper wasps (Polistes dominula) have larger poison glands". Frontiers in Zoology. 9 (1): 20. doi:10.1186/1742-9994-9-20. ISSN 1742-9994. PMC 3495029. PMID 22901602.
External links
- National Geographic News: Wasps Can Recognize Faces. Retrieved 2012-12-04.