Liostenogaster vechti

Last updated

Liostenogaster vechti
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Genus: Liostenogaster
Species:
L. vechti
Binomial name
Liostenogaster vechti
(Turillazzi, 1988)

Liostenogaster vechti is a type of eusocial hover wasp within the family Vespidae. [1] They are typically brown and yellow in color and are considered a passive aggressive species. [2] Their stings are less painful to humans than other social wasps, and they engage in associative nest foundation. They are mostly found on the Malaysian peninsula and are known for living in large clusters of small ring-shaped nests. [1]

Contents

Young colony of Liostenogaster vechti. Peninsular Malaysia Liostenogaster Vechti Photo. David Baracchi.jpg
Young colony of Liostenogaster vechti. Peninsular Malaysia

Taxonomy and phylogeny

Liostenogaster vechti was first classified by Stefano Turillazzi in 1988. These wasps are part of the subfamily Stenogastrinae and are often referred to as hover wasps due to their manner of flying. Stenogastrinae has many other groups that are distributed across the Oriental region, but the species most closely associated with L. vechti is L. flavolineata due to similar coloring, behavioral patterns, and the similar locations of the two species. [3]

Description and identification

The adult females are 14 to 16 mm in length with dark brown coloring and yellow markings all over the body. The abdomen has brown and yellow stripes with the tip being mostly dark brown, and the legs are typically lighter at the bottom and browner near the top. Their mandibles are also yellow and each have three teeth (tridentate) which are not yellow. The antennae are partially dark brown and the eyes are dark brown and hairless unlike the hairy eyes of L. flavolineata. The top of the thorax is usually dark brown with the rest of the thorax having yellow and brown intermediate patches, and the petiole (slender stalk connecting the abdomen to the thorax) is usually a reddish brown. [1] They also have two sets of wings, two posterior and anterior. [4]

The adult males look very similar to the females. They have about the same body length, and the main differences include their antennae being completely dark brown, and having hair on the broad yellow plate (clypeus) at the front of their heads. They have more hair in other places as well. Also, their upper hind limbs (femora) on their median legs are brown with a thin, long, yellow stripe. Lastly, the aedeagus (reproductive organ which is used to secrete sperm) is flattened and inflated at the apex. [1]

Habitat and distribution

Along with most other members of the Stenogastrinae, L. vechti are only located in Southeast Asian rainforests. These wasps along with L. flavolineata are mostly distributed in the Malaysian peninsula. Their nests are located in large clusters and are usually found on flat, vertical, or horizontal substrates. [5]

Colony cycle

Colonies are often founded by a group of L. vechti females using what is known as associative foundation. [6] They begin by constructing small nests in the center, and eventually other wasps join the colony and build nests around the initial central one. [3] The nests last for multiple years, and colonies grow and contain up to 600 nests or more. [7] Temperature does not play a serious limiting factor in the lives of these wasps, and there is no particular season during which the colonies of these wasps are not present and active. [6] However, brood rearing and reproduction is more likely to occur during the dry season, and the population of the nest tends to increase during that time as well, although brood rearing and reproduction also occurs during the wet season. [6]

Nests are sometimes destroyed by predators or by other environmental factors such as floods and earth slides, but new nests are continually built around the old nests. [6] Each nest usually contains relatively few individuals (usually no more than 10) and there are usually more females than males. There is typically one dominant reproductive female per nest, and after her daughters mature, they stay with their mother in her nest for a short while and engage in nest defense, brood care, or foraging activities. After this period ends, daughters often leave and join other colonies. [8] Occasionally, dominant females are usurped by other visiting dominants. A new reproductive female may arrive, fight the current dominant female, and take over the nest if she (the newcomer) wins. [3]

Nests

Nest building

L. vechti build their nests in sheltered areas on flat, vertical, or horizontal surfaces (such as on the faces of overhanging rocks). They begin by building cells one next to the other in a row by using bits of vegetation. Once they have finished the initial row, they will often build a second row atop the first one [1] and possibly even a third row if conditions are especially crowded. [5] The wasps occasionally build nests that form a complete ring which have no more than 40 cells in one row, [5] but usually the nests resemble an open ring with the open cells facing the center. This has caused the nests to be characterized as "bracket-like" or "horseshoe-like." The cells also face slightly downward to prevent raindrops from landing inside of the nests, and the nests usually contain a couple of protruding points which the wasps use as locations upon which to land. Most nests are small, but the nests are built in clusters that range from only having a few nests to containing hundreds. [1]

Paper nest of aL. vechti colony. Peninsular Malaysia Liostenogaster Vechti. Photo. David Baracchi.jpg
Paper nest of aL. vechti colony. Peninsular Malaysia

Nests are also built to take advantage of camouflage and concealment techniques to avoid predation. Old nests that are no longer used are not completely torn down or abandoned but combined with other functioning nests to form a pattern that often contributes to predator confusion. [5]

Colony behavior

At sunrise, the population of wasps currently on the nest drops to around 60%. This percentage remains about the same throughout the day as different wasps return and leave for various reasons, and the number increases to about 90% as night falls. However, the number never reaches 100% because some individuals do not return during the night. [3]

When on the nest, the majority of the wasps spend their time resting. This posture usually includes holding the abdomen to one side, and individuals will often lie on top of each other while resting. Most activity seems to be concentrated in the middle of the day, including egg-laying and abdominal pap collection. [3]

Throughout the day, some wasps return to their own nests, while others appear to "visit" or "inspect" other nests. They do this by either approaching other nests, touching other nests, or directly landing on the nests. Responses to these visitors by wasps that actually reside on these nests range from ignoring the newcomers, accepting the newcomers, or threatening and attacking the intruders. Falling fights and other more aggressive defensive maneuvers are more frequent on nests located in the center of the clusters. This suggests that while all the nests are defended, central nests are defended more aggressively, even against visitors. [5]

Spatial arrangement

Clusters are often built in an oval shape starting from the center. The center is the most coveted space, so the nests tend to be more concentrated and often smaller and older in these areas. These central nests are also more likely to have completely closed rings rather than open rings. Many nests on the periphery are larger, younger, and contain more cells, and foundations are also mainly found near the peripheries. Also, because clusters last for more than one year and some nests are destroyed during that time, signs of older nests are also more concentrated near the middle. [5]

Costs and benefits of group living

Unlike wasps in the subfamily Polistinae, Stenogastrinae are unable to construct peduncles for their nests. Being forced to choose sites with already made suspensions severely limits their options for suitable nesting sites. This also provides a reasonable explanation for why wasps belonging to this family tend to nest in clusters. When one suitable site is located, many wasps can benefit from the same site. [5]

However, there are obvious costs to group living as well. The larger the cluster, the more visible the cluster becomes and the more obvious it is to predators. While the nests do tend to blend in well with the surrounding environment, predators notice the nests, return repeatedly to the same clusters, and use them as a ready food source. [5]

Also, the nests located near the periphery are more likely to be attacked than the ones in the center which are usually completely surrounded by other nests. This is an example of the selfish herd theory and demonstrates how individuals may cluster together for protection against predators, but the ones in the middle tend to be safer than those on the outside. However, each colony is more prone to attack if the colony next to it is attacked. This is an example of the attacked neighbor risk and reduces some of the benefit gained by participating in selfish herd tactics. [5]

Reproduction

Egg-laying

As is characteristic of Stenogastrinae, L. vechti secrete a type of gelatinous substance (which is produced by the Dufour’s gland) from their abdomens, which they use to care for the offspring and to lay their eggs. The females collect the pap with their front legs and then hold it in their mouths while they lay eggs, and pap collection is performed before and after each egg is laid. [3]

Egg-laying is done by bringing the abdomen close to the mouthparts where the pap is being held. The female holds her front legs up and open and appears to use them to touch her abdomen. Her antennae are completely still and her wings are closed during the operation. Then her abdomen begins to vibrate, and her sting is turned inside and out and folded backwards. During this process, the egg is expelled and placed directly on the pap that the female is holding in her mouth. She does not touch it with her legs at all. Then her abdomen relaxes and the female places the egg inside the cell. The average length of the eggs laid by L. vechti is 1.36 mm, and ovarian eggs are considered ready to be laid once they reach 1.16 mm. [3]

Development

L. vechti go through four larval instars (developmental stages), with possibly a 5th initial instar, [4] and the total developmental period lasts over 100 days. As the larvae mature, they develop salivary glands, and afterwards they enter a pupal stage which can last for over 36 days. [4] After individuals emerge from their cells, they spend a few days residing on their nests before engaging in normal nest behavior such as nest defense, or before becoming foragers and retrieving food for the colony. [5]

Genetic relatedness

Like most hover wasps, L. vechti reproductive females mate singly and are therefore more closely related to the brood than any of the other nestmates are. This is especially true considering that most of the daughters leave the nest, and the current nestmates are often not related to the queen. [6]

Behavior

Feeding

L. vechti frequently request trophallaxis from other members within their colonies. This behavior is most exhibited whenever a forager returns to the colony. A fellow colony member may solicit food from the individual in a variety of ways. The solicitor may approach the forager and bite her on her clypeus or her mouthparts. The two wasps may also push their heads against each other as if they are fighting. While this happens, they either do not move their antennae or hold them quivering to one side, and their abdomens are pointed downwards slightly. A wasp might also solicit food by rubbing the sides of the other individual's head with her front legs and flapping her wings vigorously. Trophallaxis usually happens too quickly to be observed, and a forager will often leave the nest a few seconds after arriving because she is often rushed by many wasps at the same time. [3] Foragers also often return with insects (such as ants or termites) [6] which they have chewed up and feed to the larvae. [3]

Signaling

Nestmate recognition and the establishment of reproductive status and dominance are often maintained through facial markings and chemical signaling. [7] The bodies of L. vechti are covered by a mixture of cuticular lipids and long hydrocarbons. Females with ovarian development have much more of a particular hydrocarbon on their cuticles than females with undeveloped ovaries. This hydrocarbon appears to be a type of fertility signal which fertilized females use to show their status. [8]

The size of the brown facial markings on the heads of females also serves as a visual cue for members of the colony and serves as a badge of higher ranking status and reproductive status. Females with larger brown facial markings also have larger ovaries. These visual cues help other wasps to not only determine if a wasp is of the same species, but also to determine what to do and how to distribute themselves around the nests. Visiting females that lack these brown visual marks are more likely to be accepted into foreign nests. Acts of aggressions are much more likely to be exhibited towards a female with these distinctive markings in order to maintain a control system within the nest. [7]

In the foreground of the photo, you can appreciate the difference in the size of the facial marking of three females. Liostenogaster Vechti Photo Baracchi David.jpg
In the foreground of the photo, you can appreciate the difference in the size of the facial marking of three females.

Division of labor

Each individual nest usually contains one dominant female with well-developed ovaries. This female spends the majority of time on the nest and engages in egg-laying and abdominal pap collection. Other females act as foragers and spend most of their time away from the nest. These females are usually not fertilized and have very poor ovarian development. [3]

Dominance

The most acts of dominance are usually performed by females with the highest ovarian indices. They will sometimes bite a subordinate’s face and sometimes flap their wings. The subordinate will often lower its head, cease voluntary movement, and sometimes even release a drop of liquid from its abdomen as a deferential act. The dominant wasp ceases movement of her own antennae and may or may not accept the drop. [3]

Fighting

When disputes occur within the nests, individuals may fight each other to determine dominance. Two wasps may engage in this activity by holding their antennae out-stretched, grasping each other, and falling from the nest. The wasps never actually hit the ground. Instead they separate in mid-air, return to the nest, and repeat this process until a winner is determined. This behavior is especially likely in the event that a stranger attempts to take over a nest while the dominant female is away. When the dominant female returns, she will "fight" this individual, often for several hours and possibly even a couple days, to determine who will be in charge of the nest. [3]

Predators and defense

When an individual of L. vechti feels threatened, it will assume a menacing stance by bending its abdomen to one side and pointing the tip in the direction of its challenger. It does this so that it is prepared to sting its opponent if necessary and thrust its abdomen toward the intruder in what is known as an abdominal slap. [3]

The main predators of these wasps are hornets (especially Vespa tropica ), ichneumonid flies, and ants. [5] The hornets attack and damage the nests by ripping the cells apart with their mandibles and then extracting portions of the brood. Raided nests can easily be identified by locating cells that have been ripped or torn. [5] When a hornet or other flying predator approaches, the wasps engage in aerial attacks by throwing themselves into the air in the direction of the potential danger. They actually engage in this behavior very frequently and do so in response to any insect of similar size to hornets that approaches the nests in a questionable manner. When ants approach or invade the nests, the wasps will either grab the ants and drag them away from the nests, or, if the ants are especially large, the wasps will first assume a defensive posture, seize the ants, and then purposely fall out of the nests to ensure the ants are removed. [3] Spiders may also prey on these wasps, but they are much more likely to catch solitary individuals away from the nest. [3]

Venom

Although they have effective stingers, these wasps and other members of Stenogastrinae are typically passive aggressive. Their stings are less painful for humans than those of many other social wasps, and often when faced with a large threat, these wasps will simply fall passively out of their nests. This behavior actually makes it harder for the potential predator to determine the location of the nest. [2]

Related Research Articles

<span class="mw-page-title-main">Yellowjacket</span> Common name for two genera of wasps

Yellowjacket or yellow jacket is the common name in North America for predatory social wasps of the genera Vespula and Dolichovespula. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of these are black and yellow like the eastern yellowjacket and the aerial yellowjacket ; some are black and white like the bald-faced hornet. Others may have the abdomen background color red instead of black. They can be identified by their distinctive markings, their occurrence only in colonies, and a characteristic, rapid, side-to-side flight pattern prior to landing. All females are capable of stinging. Yellowjackets are important predators of pest insects.

<span class="mw-page-title-main">Paper wasp</span> Vespid wasps that gather fibers from dead wood and plant stems

Paper wasps are vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrella wasps, due to the distinctive design of their nests.

<span class="mw-page-title-main">European paper wasp</span> Species of wasp

The European paper wasp is one of the most common and well-known species of social wasps in the genus Polistes. Its diet is more diverse than those of most Polistes species—many genera of insects versus mainly caterpillars in other Polistes—giving it superior survivability compared to other wasp species during a shortage of resources.

<span class="mw-page-title-main">Stenogastrinae</span> Subfamily of wasps

The Stenogastrinae are a subfamily of social wasps included in the family Vespidae. They are sometimes called hover wasps owing to the particular hovering flight of some species. Their morphology and biology present interesting peculiarities.

<i>Polistes fuscatus</i> Species of insect

Polistes fuscatus, whose common name is the dark or northern paper wasp, is widely found in eastern North America, from southern Canada through the southern United States. It often nests around human development. However, it greatly prefers areas in which wood is readily available for use as nest material, therefore they are also found near and in woodlands and savannas. P. fuscatus is a social wasp that is part of a complex society based around a single dominant foundress along with other cofoundresses and a dominance hierarchy.

Belonogaster juncea juncea is a subspecies of Belonogaster juncea and is classified as a primitively eusocial wasp, meaning that the species is social while exhibiting a morphology that is indistinguishable from that of other castes. It is also classified as a type of African Paper Wasp. Many of the studies relating specifically to B. j. juncea take place at the University of Yaoundé in Cameroon.

<i>Polistes nimpha</i> Species of wasp

Polistes nimpha is a eusocial paper wasp found all over Europe, with particular sightings in Turkey, Finland, Estonia, and Latvia. It is also found in northern Africa, Pakistan, Iran, India, Kazakhstan, Mongolia, and China. The climate in these areas is relatively cold and snowy in the winter, while summers are usually hot and dry, with steppe vegetation. Polistes nimpha colonies are relatively small and easily manipulated.

<i>Liostenogaster flavolineata</i> Species of wasp

Liostenogaster flavolineata is an insect that belongs to the wasp family Vespidae. This hairy-faced hover wasp species is predominantly found in South Asian rain forests, especially in Malaysia. Individual colonies of this species are very small, but aggregations of nests allow for interactions between many smaller colonies. Some worker wasps, known as "helpers", will move between multiple nests in an attempt to improve their position in the dominance hierarchy. Its nests are pale-colored and are usually built with mud. Liostenogaster flavolineata is one of the most studied species in the Stenogastrinae.

<i>Parischnogaster mellyi</i> Species of wasp

Parischnogaster mellyi is a medium-sized species of a hover wasp in the family Vespidae. It is found in Southeast Asia and is widely spread in Thailand and Malaysia. Its nests feature flexible and dynamic qualities, and they are commonly seen under roofs of houses and huts in rural areas. Hovering and patrolling behaviors are the species’ main defining behavioral features, and such activities are closely linked to its mating patterns.

<i>Polistes semenowi</i> Species of wasp

Polistes semenowi is a species of paper wasp in the genus Polistes that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name Polistes sulcifer, while a different species was incorrectly believed to represent P. semenowi. It is one of only four known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. semenowi females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.

<i>Ropalidia revolutionalis</i> Species of wasp

Ropalidia revolutionalis, the stick-nest brown paper wasp, is a diurnal social wasp of the family Vespidae. They are known for the distinctive combs they make for their nests, and they are mostly found in Queensland, Australia in the areas of Brisbane and Townsville. They are an independent founding wasp species, and they build new nests each spring. They can be helpful because they control insect pests in gardens.

<i>Parischnogaster alternata</i> Species of wasp

The Black hover wasp, Parischnogaster alternata, is a eusocial wasp in the genus Parischnogaster. It is native to South-East Asia, and builds its nests in cavities located in dark and damp locations. The nests of Black hover wasps are often found in clusters, which serves as a passive defense mechanism against predators. The annual colony cycle begins with nest initiation by a single foundress though colonies typically consist of 2-3 associative females and helpers that aid in brood development, nest construction, and colony defense. Indicative of the name, female P. alternata are known to strategically hover near nests when visiting other colonies before landing. These intrusions produce responses ranging from aerial fighting to cooperative food sharing.

Parischnogaster nigricans serrei is a hover wasp subspecies in the family Vespidae, and it is predominantly found in the Java region of Indonesia. Its nest cells are of conical structure, linearly attached to a string-like substratum. The nests are typically found in places open to human interactions, such as gardens, trees, or forests around villages. There is a clear dominance hierarchy within colonies, which often affects the behavioral activities of its members. The wasp’s most common predators are Vespa tropica, also known as the great banded hornet. P. nigricans serrei defends itself by flying away or giving out alarm calls.

<i>Polistes japonicus</i> Species of wasp

Polistes japonicus is a eusocial paper wasp found in Japan. It was first described by Henri Louis Frédéric de Saussure in 1858. It is closely related to Polistes formosanus. This species lives in small colonies with few workers and a foundress queen. Nests of these wasps are sometimes used as a traditional medicine in Korea, China, and Japan.

<i>Mischocyttarus flavitarsis</i> Species of wasp

Mischocyttarus flavitarsis is a social paper wasp found in western North America. Their nests can be found both in forests close to rivers or in close proximity to human life under the eaves of roofs. Despite the fact that M. flavitarsis nests are frequently in close contact with humans, M. flavitarsis typically will not sting, but rather ram into the threatening individual. Their colony cycle typically begins before May and will last until October. The queen will then seek a hibernation site for the winter. Perched near female hibernation sites are males with whom the female will mate. The males have claimed their territory by rubbing sternal brushes along the border of the site, leaving a chemical that deters other individuals from approaching. M. flavitarsis feed on arthropods, nectar, and animal carcasses and are often prey to birds, ants, and praying mantis.

Parachartergus fraternus is a neotropical, swarm founding, polistine wasp species that is distributed throughout Central and South America. They live in nests in second growth tropical dry forests, near pasture fields, roadside areas, and urban areas as well. These wasps eat insects, such as caterpillars of Lepidoptera. They also drink nectar, honeydew, and water. The workers capture their prey during foraging. They also use venom to paralyze their prey in order to consume it later. P. fraternus wasps are not very aggressive and they do not attack when the nest is approached.

<i>Parischnogaster striatula</i> Species of wasp

Parischnogaster striatula is a species of social hover wasps found in Southeast Asia. Their nests are uniquely shaped, mimic their surroundings and, like the other Stenogastrinae social wasps, lack a nest pedicel. They are also unique in their use of glandular secretion when laying eggs. P. striatula is also different than typical wasps because the queens are not the only ones who possess reproductive capabilities; other females also have developed ovaries. This wasp is also known for fiercely protecting its nest against enemies, such as ants or other hornets.

Polistes erythrocephalus is a species of paper wasp in the subfamily Polistinae of family Vespidae found in Central and South America. P. erythrocephalus is a eusocial wasp, meaning that it possesses both reproductive and non-reproductive castes. The cooperation between the two castes to raise young demonstrates the altruistic nature of these wasps. P. erythrocephalus exhibits a four-stage colony cycle, as do many other Polistes wasps. This species generally feeds on larvae, occasionally their own, and is preyed upon by species such as army ants.

<i>Parischnogaster jacobsoni</i> Species of wasp

Parischnogaster jacobsoni is a species of social wasp within Parischnogaster, the largest and least known genus of Stenogastrinae. It is distinguished mainly by its tendency to construct ant guards on its nests. Natural selection has led this wasp to have a thick substance emitted from its abdominal glands that allows it to protect its nest from invasions. Parischnogaster as a genus has been relatively unstudied; P. jacobsoni is one of the few investigated species because it has sufficient durability to live near human populations and it has demonstrated unusual resilience to pollution. While P. jacobsoni is a more complex organism than other wasps in Parischnogaster, the genus overall is relatively primitive with respect to social wasps as a whole.

<i>Liostenogaster</i> Genus of wasps

Liostenogaster is a genus of hover wasps from the subfamily Stenogastrinae of the family Vespidae which has a distribution centred on south-east Asia. It was named by the Dutch entomologist Jacobus van der Vecht from material collected by Japanese scientists on an expedition to Thailand, Malaysia and Cambodia which took place in 1966.

References

  1. 1 2 3 4 5 6 Turillazzi, Stefano (1988). "Adults and nest of Liostenogaster vechti n. sp. (Hymenoptera Stenogastrinae)". Tropical Zoology. 1 (2): 193–201. doi: 10.1080/03946975.1988.10539414 .
  2. 1 2 Francesca R. Dani; E. David Morgan; Graeme R. Jones; Stefano Turillazzi; Rita Cervo; Wittko Francke (1998). "Species-Specific Volatile Substances in the Venom Sac of Hover Wasps". Journal of Chemical Ecology. 24 (6): 1091–1104. doi:10.1023/A:1022358604352. S2CID   35475771.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Turillazzi, Stefano (1990). "Social biology of Liostenogaster vechti Turillazzi 1988 (Hymenoptera Stenogastrinae)". Tropical Zoology. 3 (1): 69–87. doi: 10.1080/03946975.1990.10539449 .
  4. 1 2 3 Turillazzi, Stefano (2012). The Biology of Hover Wasps. Springer Berlin Heidelberg. pp. 27–60.
  5. 1 2 3 4 5 6 7 8 9 10 11 12 Coster-Longman, Landi, and Turillazzi, C.,M., and S. (2002). "The Role of Passive Defense (Selfish Herd and Dilution Effect) in the Gregarious Nesting of Liostenogaster Wasps (Vespidae, Hymenoptera, Stenogastrinae)". Journal of Insect Behavior. 15 (3): 331–350. doi:10.1023/A:1016213125161. S2CID   11563430.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. 1 2 3 4 5 6 Turillazzi, Stefano. The Biology of Hover Wasps. Springer Berlin Heidelberg. pp. 89–127.
  7. 1 2 3 Baracchi D.; Petrocelli I.; Cusseau G.; Pizzocaro L.; Teseo S.; Turillazzi S. (2013). "Animal Behaviour: Facial markings in the hover wasps: quality signals and familiar recognition cues in two species of Stenogastrinae". Animal Behaviour. 85: Volume 85 Issue 1 203–212. doi:10.1016/j.anbehav.2012.10.027. hdl: 2158/790326 . S2CID   53203615.
  8. 1 2 Turillazzi, S.; Sledge, M. F.; Dapporto, L.; Landi, M.; Fanelli, D.; Fondelli, L.; Zanetti, P.; Dani, F. R. (2004). "Epicuticular lipids and fertility in primitively social wasps (Hymenoptera Stenogastrinae)". Physiological Entomology. 29 (5): 464–471. doi:10.1111/j.0307-6962.2004.00418.x. S2CID   86685690.