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''Vespula squamosa'', the southern yellowjacket, is a
social 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. Th ...
. This species can be identified by its distinctive black and yellow patterning and orange queen. These yellowjackets are typically found in eastern
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, and its territory extends as far south as
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
. Within these territories, they create enormous, multiple-comb nests.MacDonald, John F., Matthews, R. W. "Nesting Biology of the Southern Yellowjacket, ''Vespula squamosa'' (Hymenoptera: Vespidae): Social Parasitism and Independent Founding", "
Journal of the Kansas Entomological Society The ''Journal of the Kansas Entomological Society'' is a quarterly peer-reviewed scientific journal published by the Kansas Entomological Society. The journal has a 2009 impact factor The impact factor (IF) or journal impact factor (JIF) of an ...
", January 1984.
The colonies may be either annual or perennial depending on the climate, and in many perennial nests, polygyny takes place. In addition, this species uses pheromones both as a sexual attractant and an alarm signal.Landoldt, P. J. and Heath, R. R. "Alarm Pheromone Behavior of Vespula Squamosa (Hymenoptera: Vespidae)", "
Florida Entomologist The ''Florida Entomologist'' is an quarterly open access scientific journal published by the Florida Entomological Society. Founded in 1917 as “The Florida Buggist” and in 1920 was renamed, into “The Florida Entomologist.” Manuscripts from ...
". June 1987.
This species feeds on insects and animal carcasses; it does not produce honey. ''V. squamosa'', a social insect, has developed a parasitic relationship with the species ''V. vidua'' and '' V. maculifrons.'' Due to their painful, venomous stings, the species is considered a pest.


Taxonomy and phylogeny

''V. squamosa'' is commonly called the southern yellowjacket. This species was discovered by
Dru Drury Dru Drury (4 February 1724 – 15 December 1803) was a British collector of natural history specimens and an entomologist. He had specimens collected from across the world through a network of ship's officers and collectors including Henry Sme ...
around 1770. It is a member of the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
, which includes paper wasps (subfamily
Polistinae The Polistinae is a subfamily of eusocial wasps belonging to the Family (biology), family Vespidae. They are closely related to the more familiar wasps (“yellowjackets” as they are called in North America) and true hornets of the subfamily V ...
), pollen wasps (subfamily
Masarinae Pollen wasps, the Masarinae, are unusual wasps that are typically treated as a subfamily of Vespidae, but have in the past sometimes been recognized as a separate family, "Masaridae", which also included the subfamilies Euparagiinae and Gayellina ...
), potter and mason wasps (subfamily
Eumeninae Potter wasps (or mason wasps), the Eumeninae, are a cosmopolitan wasp group presently treated as a subfamily of Vespidae, but sometimes recognized in the past as a separate family, Eumenidae. Recognition Most eumenine species are black or brown, ...
), and
yellow jacket Yellowjacket or yellowjacket 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 ...
s (subfamily
Vespinae The subfamily Vespinae contains the largest and best-known eusocial wasps, including true hornets (the genus ''Vespa''), and the "yellowjackets" (genera ''Dolichovespula'' and ''Vespula''). The remaining genus, '' Provespa'', is a small, poorly ...
). The family Vespidae has about 4000 species, the majority of which are not social. Within the subfamily Vespinae, however, the roughly 80 species are all eusocial, including ''V. squamosa''. They create either annual or perennial nests, and some species, including ''V. squamosa'', are
parasites Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
to other Vespinae species. Within the subfamily Vespinae, the four genera are ''
Provespa ''Provespa'' is a small genus of Vespidae, made up of nocturnal wasps from Southeast Asia, sometimes referred to as "night wasps" or "night hornets", though they are not true hornets (genus ''Vespa''). They are the only nocturnal members of the ...
,
Vespa Vespa () is an Italian luxury brand of scooter (motorcycle), scooters and mopeds manufactured by Piaggio. The name means wasp in Italian. The Vespa has evolved from a single model motor scooter manufactured in 1946 by Piaggio & Co. S.p.A. of ...
,
Dolichovespula ''Dolichovespula'' is a small genus of social wasps distributed widely throughout the Northern Hemisphere. The yellow and black members of the genus are known by the common name yellowjackets in North America, such as ''Dolichovespula norwegica'' ...
'', and ''
Vespula ''Vespula'' is a small genus of social wasps, widely distributed in the Northern Hemisphere. Along with members of their sister genus ''Dolichovespula'', they are collectively known by the common name yellowjackets (or yellow jackets) in North ...
''. Within the genus ''Vespula'' are 18 species.Kimsey LS, Carpenter JM (2012) "The Vespinae of North America (Vespidae, Hymenoptera)","
Journal of Hymenoptera Research The ''Journal of Hymenoptera Research'' is a peer-reviewed scientific journal covering systematics, taxonomy, and ecology of Hymenoptera. It was established in 1992, and transferred to publishing with Pensoft Publishers in 2011, under an open acces ...
"
The southern yellowjacket is fairly closely related to ''V. maculifrons'' (eastern yellowjacket), and ''V. squamosa'' is a social parasite of ''V. maculifrons''. ''V. squamosa'' is also closely related to '' V. rufa''.


Description and identification

''V. squamosa'' is typically about long and distinguishable by its black body and yellow striping pattern over its entire body. This species has clear wings and a hairless body. While the males and workers resemble other yellowjackets, the queen has a unique appearance. She is significantly larger than the males and workers and is more orange in color."The Yellowjackets of American North of Mexico". Retrieved on 21 September 2014.
/ref> In addition, both ''V. squamosa'' and the ''V. sulphurea'' have two stripes on their scuta, which differentiates them from other species.


Distribution and habitat

Southern yellowjackets are typically found in the eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
and as far south as
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by H ...
. Their territory expands as far west as
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
and as far east as the
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
. These areas tend to have fairly warm weather and in some cases, tropical climates. Within these territories, nests are typically found in unnatural habitats, such as yards, parks, and sides of roads. Some colonies, though, are found in pine forests. Their parasitic relationship with ''V. maculifrons'' can result in ''V. squamosa'' colony formation in hardwood forests, a habitat more typical of ''V. maculifrons''. Most nests are
subterranean Subterranean(s) or The Subterranean(s) may refer to: * Subterranea (geography), underground structures, both natural and man-made Literature * ''Subterranean'' (novel), a 1998 novel by James Rollins * ''Subterranean Magazine'', an American fa ...
, though some have been found above ground or in walls, close to ''V. maculifrons'' nests.


Nests

Southern yellowjackets' nests are typically built in unnatural or disturbed environments such as yards, picnic tables, and other artificial environments. They are usually made from chewed vegetable fibers. Colonies have been reported to contain from 9000 to more than 450,000 cells. One of the largest reported nests was in diameter at its widest point and had about 39 combs. It was about tall and had the same diameter. While most social wasps have annual nests, many instances of multiple-season ''V. squamosa'' nests have been found, especially in the southern coastal areas of the species' range.Pickett, K. M., Osborne, D. M., Wahl, D., and Wenzel, J. W. (2001). "An Enormous Nest of Vespula squamosa from Florida, the Largest Social Was Nest Reported from North America, with Notes on Colony Cycle and Reproduction", "
Journal of the New York Entomological Society The New York Entomological Society and other entomological societies in New York have produced a number of scientific journals since the mid-19th century, some of which have moved between a set of similar societies. ''Bulletin of the Brooklyn Ent ...
", 2001


Colony cycle


Annual colonies

The colony cycle for southern yellowjackets is fairly variable. Many annual colonies begin in late May or early June when the queen infringes on a small ''V. maculifrons'' colony (about 85% of colonies begin in this parasitic takeover, while only 15% of colonies are independent). Whereas workers do not survive the winter in annual colonies, the queen does survive, and in the spring, when she is ready to lay more eggs, she scouts for small ''V. maculifrons'' and ''V. vidua'' colonies to serve as the host colony. These “takeovers” can often be violent as evidenced by many dead queens found in the nests. This violence tends to be
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...
, meaning that multiple ''V. squamosa'' queens are fighting for control of the host nest or for control of the
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
nest. A successful usurpation constitutes the beginning of the parasitic relationship between a ''V. squamosa'' colony and a ''V. maculifrons'' colony. These colonies are built around the host nest, thus increase the size of the initial host nest. The queen begins the building process by constructing queen cells and laying eggs that will hatch
larvae 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. The ...
. The larvae enter the pupal phase after about a month. Once more of these female workers have been produced (late July or early August), the colony size begins to grow significantly as worker cell construction begins. This period from July to August is the major growth period during which up to 4000 workers could be producing cells. Workers build smaller 4.0- to 5.0-mm-wide cells, while the queens build “queen cells” which are about 7.0 mm wide. Most nests have their maximum number of queen and worker combs between October and December, the end of the seasonal colony cycle. As the season ends, the queen hibernates so she can, in many cases, begin a new colony the next year.


Overwintering colonies

While most colonies are annual,
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
colonies exist, especially in
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. These nests tend to be found in the southernmost regions of their habitat where it is possible for the nest to survive through the winter. Because these nests are built for and inhabited for several years, they tend to be much larger than annual nests. In fact, these nests are about 50-60 liters in volume and can contain at least 60,000 cells. The decreased production of the winter brood further distinguishes overwintering nests from annual nests. In addition, continuous production of reproductives occurs even through the winter. These nests also have from six to over 100 reproductive queens.


Mating behavior


Mating system

Southern yellowjackets have a
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
mating system: a colony has more than one queen. In other''
Vespula ''Vespula'' is a small genus of social wasps, widely distributed in the Northern Hemisphere. Along with members of their sister genus ''Dolichovespula'', they are collectively known by the common name yellowjackets (or yellow jackets) in North ...
'' species, polygyny tends to be found in mild climates where the colony can last more than a year. Given that many of this species live in temperate or subtropical environments, perennial nests have been observed. Polygynous perennial colonies occur when reproductive females (queens) reproduce in the colony instead of leaving it. As a result of the multiple queens and additional seasons and workers in perennial nests, these colonies tend to be much larger than their annual counterparts. While scientists have argued that a polygynous mating system would breed disease and reduce fitness, many examples of the polygynous nests are seen. The parasitic nature of the species goes hand in hand with their polygynous mating system. In a parasitic system, females lay eggs in a different species’ nests, and in a polygynous system a female of the same species lays eggs in the nest in the hopes of overtaking the current queen. Dead queens of the same species have been found in ''V. squamosa'' nests, supporting the idea of competition between queens as a result of polygyny. This polygyny may be tolerated in large multiple-season nests because the size of the nest reduces interactions between the competing queens. This process of seizing the nest within the species could provide the basis for parasitic nature of this species, as the takeover of a nest either within species or between species is a fairly similar process.


Mating of the queen

Just as workers are able to communicate via
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
to co-ordinate attacks, the queen also produces her own pheromones. Whereas the workers had a pheromone to spread alarm, she has a pheromone to attract male mates. Being the sole reproductive female in the nest, the queen must mate, so these attraction pheromones are very important. Thus, her job is to successfully mate to populate a new nest with workers, males, and females in a new nest the next season.


Sexual dimorphism

Sexual and caste dimorphism is very pronounced in this species. While all individuals have a distinctive marking on their scuta, workers, males, and queens all have different appearances. Workers and males have similar black and yellow striped patterning, but the queen has fewer black markings and a more orange/brown coloring. Wing length also helps to differentiate between the worker and male. Workers have fore wing lengths of 9.5-11.0 mm, and males’ are about 12.5 mm. In addition, the queen is about 4 mm longer than her worker and male counterparts.Buck, Matthias et al

'' Canadian Journal of Arthropod Identification'', February 2008. Retrieved 22 September 2014.


Defense and alarm signals

As a form of defense of their nests, southern yellowjacket workers use alarm
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
to communicate with each other to coordinate an attack. These behaviors are chemically mediated, and these alarm pheromones cause many social wasp species to leave the nest and attack whatever may be threatening it. Individuals can emit this pheromone from a few places in their bodies. Pheromone activity occurs in the stomach, and more specifically in a venom gland within the stomach. Odors on objects or people attacked by ''V. squamosa'' differ from the previously isolated chemical alarm pheromone, N-3-methylbutylacetamide. A second source of an alarm pheromone is found in the venom glands of the head of an individual. The alarm pheromones may be applied by the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
.Landoldt, P. J., Reed, H. C., and Heath, R. R. "An Alarm Pheromone from Heads of Worker Vespula squamosa (Hymenoptera: Vespidae)", "
Florida Entomologist The ''Florida Entomologist'' is an quarterly open access scientific journal published by the Florida Entomological Society. Founded in 1917 as “The Florida Buggist” and in 1920 was renamed, into “The Florida Entomologist.” Manuscripts from ...
", June 1999.


Diet

This species is predatory and typically eats live insects, but they also feed on the flesh of deceased prey. They typically prey on
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
, including
spiders 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 dive ...
and
caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symph ...
. For
larvae 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. The ...
to grow into the pupal state, adult workers find prey and bring food back for them. When establishing the colony, the queen goes out in search of
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and insects for the larvae, as well. This species does not produce honey, though. "Southern Yellowjacket" Buck, Matthias et al

''Canadian Journal of Arthropod Identification'', February 2008. Retrieved 22 September 2014.


Interactions with other organisms


Parasitic relationships

Most southern yellowjackets are parasitic, in that they benefit at the expense of another species, as seen in their nest formation, as they build their nests around those of another species. Only about 15% of the nests are independently founded. Evidence shows ''V. squamosa'' as a social parasite of ''V. vidua'' (ground hornet) and ''V. maculifrons'' (
eastern yellowjacket The eastern yellow jacket or eastern yellowjacket (''Vespula maculifrons'') is a wasp found in eastern North America.Hoffman, Eric A., Kovacs, Jennifer L. and Goodisman, Michael A. D. (August 20, 2008). Genetic structure and breeding system in a ...
). As this species depends on other species for part of its nest, the location of its nests is often contingent on the location of the host's nests.


Human interactions

This species is currently abundant in urban areas, as their colonies tend to be found in unnatural environments (roadsides, wall voids, parks, etc.). Because of this close proximity to humans and the size of the nests, this species is typically considered a pest. The close interactions between southern yellowjackets and humans commonly lead to multiple, painful stings."The Yellowjackets of American North of Mexico". Retrieved on 21 September 2014. Their stings are venomous, which causes the pain, and workers can sting several times. These stinging attacks are typically effects of alarm and occur in defense of their colony, so disturbing a nest can result in multiple stings.Landoldt, P. J. and Heath, R. R. "Alarm Pheromone Behavior of Vespula Squamosa (Hymenoptera: Vespidae)", "
Florida Entomologist The ''Florida Entomologist'' is an quarterly open access scientific journal published by the Florida Entomological Society. Founded in 1917 as “The Florida Buggist” and in 1920 was renamed, into “The Florida Entomologist.” Manuscripts from ...
". June 1987.


References

{{Taxonbar, from=Q15119916 Vespidae Hymenoptera of North America Insects described in 1773 Taxa named by Dru Drury