Polistes Pacificus
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''Polistes pacificus'' is a Neotropical species of social
paper wasp Paper wasps are Vespidae, 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 umbrell ...
belonging to the 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 ...
and 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 ...
. ''P. pacificus'' can be found distributed throughout most of Central and South America and parts of southern North America. First discovered by
Johan Christian Fabricius Johan Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is cons ...
in 1804, ''P. pacificus'' is much darker in color than some other more recognizable ''
Polistes Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella ...
'' wasps, and is one of the insects commonly eaten by several indigenous groups in Venezuela and Colombia.


Taxonomy and phylogenetics


Phylogenetic tree

''P. pacificus'' is part of a monophyletic clade that makes up the membership of Richards’ (1973, 1978) subgenera ''Epicnemius''. ''P. pacificus'' is most closely related to its
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
, ''P. cinerascens''. Although previous studies found ''Epicnemius'' to be
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
, the most recent and complete study done by Picket et al. reveals that it is likely
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The species included in this
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
are ''P. cinerascens'', ''P. pacificus'', ''P. geminatus geminatus'', ''P. occipitalis'', ''P. testaceicolor''.


Synonyms

''P. pacificus ''was originally discovered in 1804 by the famous Danish entomologist,
Johan Christian Fabricius Johan Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is cons ...
(1745-1808). However, because ''P. pacificus'' has wide color variation depending on its geographic region, over time it has been mistaken for several new species and given new names. These names included ''P. flavipictus Ducke'', 1918 ''P. liliaceusculus de Saussure'', 1854, ''P. liliaciosus de Saussure'', 1855, ''P. modestus Smith'', 1862, ''P. trinitatis Bequard'', 1937. These names are relatively obsolete, but may appear in erroneous or outdated literature.


Description and identification


Distinguishing ''P. pacificus'' from similar species

''P. pacificus'' has varied coloration patterns with markings that can be black, ferruginous and yellow. It is darker and less brightly colored than most other ''Polistes ''species, and it is morphologically very similar to ''P. boharti.'' In fact, the two species are classified as the same under Richard's key. However, there are some distinguishing characteristics between the two: Firstly, the digitus structure of the genitalia of ''P. pacificus'' males has many dark, stout, distally curved or hooked
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
. Secondly, female ''P. pacificus'' will have a genal ridge that is usually weak below, and rarely reaches below the level of the lower one-fifth to one-fourth of the eye; on the rare occasions that it extends below this level, it will most likely not reach the base of 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 ...
and will be very weak. Additionally, in ''P. pacificus'', the lobate humeral angle is not defined, and the lower lateral extension of the pronotal ridge will not be depressed.


The defined morphological criteria

A specimen of ''P. pacificus'' may also be identified by the following morphological criteria: * Male antennae are hooked * Clypeus is pointed (see diagram) * Male eyes touch the clypeus (see diagram) * There is much pigmentation on the head * Claws are symmetrical * There are two teeth, nearly equal in size, that are present in the mandible *
Setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
on the underside of the thoracic segment are minute or short, while setae on the underside of the abdominal segment are long and hairy.


Nest identification

''P. pacificus'' builds nests that are relatively small and made of paper. These nests are most likely to be found in areas of low second-growth vegetation, and are often sheltered by large leaves. The nests, which hang from a plant supported by one or more
pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
s, are made up of a single comb of hexagonal cells and do not have an envelope. ''Polistes'' produce the water-resistant paper used to make nests by gathering wood fiber and other plant matter and combining it with saliva.


Distribution and habitat


Geographic distribution

''P. pacificus'' can be found from the southern United States to Uruguay and Argentina. ''P. pacificus'' can be found in Southern parts of Texas, various areas of Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Venezuela, Trinidad, Guyana, Suriname, French Guiana, parts of Brazil, Paraguay, Bolivia, and Uruguay.


Nest-building habitats

''P. pacificus'' builds its nest on plants, using large leaves of the host plants as shelter. While some reports have been made that suggest that social paper wasps build nests where arboreal ants are also present, ''P. pacificus'' is only rarely associated with arboreal ants and frequently chooses nesting sites on plants that are unattractive to ants. Some of the most common plants for ''P. pacificus'' to build nests on are ''
Clusia ''Clusia'' is the type genus of the plant family Clusiaceae. Comprising 300-400 species, it is native to the tropics of the Americas. The genus is named by Carl Linnaeus in honor of the botanist Carolus Clusius. The closest relatives of ''Clus ...
grandiflora'' and ''
Vismia ''Vismia'' is a genus of flowering plants in the family Hypericaceae. Members of the genus are small trees and shrubs found in tropical and subtropical areas of Central America and South America. Including the countries of Belize, Bolivia, Brazil ...
sessilifolia'', but they have also been known to build nests that connect to long, thin thorns under the leaves on juvenile ''Astrocaryum sciophylum'' palm trees.


Colony cycle

''P. pacificus'' is one of several social paper wasp species that migrate seasonally depending on the times of the wet and dry seasons. In areas with less pronounced dry seasons, ''P. pacificus'' is more likely to brood continuously. However, when ''P. pacificus'' live in regions with a defined dry season, the dry season is the non-nesting phase of the colony cycle. During this period, wasps leave the low elevation habitats in which they normally nest and reproduce during the rainy season, and move to cooler areas at higher elevation. The aggregating wasps are sometimes found to be in small groups of individuals from different nests. Due to the fact that dry season aggregates of ''P. pacificus'' have been found in evergreen old growth forests, a place distinct from where they nest, it has been suggested that this behavior is evidence of
altitudinal migration Altitudinal migration is a short-distance animal migration from lower altitudes to higher altitudes and back. Altitudinal migrants change their elevation with the seasons making this form of animal migration seasonal. Altitudinal migration can be m ...
. The benefits to this type of seasonal migration could be that wasps in these cooler high elevation conditions have lower metabolic needs and costs. This behavior is similar to migrating behavior observed in temperate zone ''Polistes'' wasps during winter. Mating can occur either at
leks A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...
away from nests or at newly founded nests, often late in the nesting season, but before the seasonal dormancy. Generally the inseminated females will pass the dormant season hibernating in a group of other females in a type of shelter, and at this time males most likely die off.


Interactions with other species


Diet

The ''Polistes'' have high nutritional demands because they nurse their broods. ''P. pacificus'' is an animal of prey, but it is also a flower-visiting wasp that gets carbohydrates from flower nectar. When ''P. pacificus'' is in the larval state, the majority of its diet comes from animal protein. In addition to the caterpillars which ''Polistes'' mostly preys on, the ''P. pacificus'' also visits flowers, thus picking up carbohydrates. Carbohydrates collected through flower nectar are a complement of its diet and thought to help maintain its own energy metabolism.


Predators

''P. pacificus'' has at least several known predators: arboreal ants and vertebrates, including humans. The larvae of the ''P. pacificus'' is what predators eat, as it is localized within the nest and immobile. The larvae makes a very valuable food source because it is rich in amino acids and fats.


Defense

''P. pacificus'' nest site selection and structure have evolved under selective pressures of weather and predation. In order to keep predators out, ''P. pacificus'' nests are built in specific places in which they are less susceptible to attack by arboreal ants or vertebrates. The placement of the nest not only aims to reduce predation, but also to provide protection from extreme weather. Nests are most often built on the underside of large leaves, which protect against weather and limit predator’s views of the nest from above. The most preferable large leaves have spines along the central vein, which act as a pedicel and protect the nests from ant attacks. In the neotropics, ''P. pacificus'' nests are most closely associated with a specific plant, the ''Clusia grandiflora'', which is most likely an adaptation due to the selective pressure of ant predation as ''Clusia'' is not an attractive host for arboreal ant nests.


Communication


Lack of visual signaling

Communication is a critical for any species and can benefit both the sender and receiver of information. Some wasps have high visual acuity and use recognition of facial patterns to establish dominance hierarchies; however, ''P. pacificus'' differs from these species in that it does not use visual signals, and must rely on other modes of communication. It is likely that ''P. polistes'' detects pheromones and other non-visual recognition cues to distinguish nestmates from nonnestmates and determine dominance.


Worker mouthing and rubbing

Adult female workers of ''P. pacificus'' have been observed to mouth and rub newly emerged female
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 ...
, during which time the newly emerged wasps remain stationary. In several cases, the older worker used her gaster to rub the new wasp with discrete anterior to posterior strokes. In other cases, the older female has also been seen to mouth the young wasps with her mandibles, by opening and closing them over the new wasp’s head,
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 cre ...
and gaster. While in contact with the younger wasp, the older worker kept her sting chamber open. The workers that were observed executing this behavior had few, small, only partially developed eggs in their ovaries, and had shriveled Dufour’s glands. Although it is unclear why exactly the wasps partook in this behavior, it is possible that doing so may give each of the newly emerged wasps the colony specific odor, which could ensure them safety when returning to the nest after foraging. It has also been hypothesized that this behavior may express some order of dominance.


Human importance


Agriculture

''P. pacificus'', which gets some of its nutrition from flower nectar, is also a
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
, and thus an important species for agriculture. In Mexico, ''P. pacificus'' has been observed to pollinate
avocado The avocado (''Persea americana'') is a medium-sized, evergreen tree in the laurel family (Lauraceae). It is native to the Americas and was first domesticated by Mesoamerican tribes more than 5,000 years ago. Then as now it was prized for i ...
s, and it has been suggested that as a native species to the area ''P. pacificus'' started a coevolution with avocado species. Although ''P. pacificus'' is not as abundant as some of the other insects known to pollinate avocados, it nevertheless contributes to avocado pollination and production.


Human food source

Not only does ''P. pacificus'' help pollinate avocados, thus helping to provide a calorie-dense plant-based food source for humans, but it also is commonly hunted by humans who eat the wasp larvae. Although insect eating is often portrayed as an archaic trait, among the Yukpa-Yuko Indians of Venezuela and Colombia, insect food sources are still vitally important, as they supplement the Yukpa’s diet by providing animal proteins and fats. The Yukpa word for ''P. pacificus'' is ''“nonawu,”'' and the Indians can recognize the nests as being low to the ground in second-growth vegetation. Although both women and men may participate in identifying nests, collecting the nests is exclusively men’s work. When it comes time to harvest, the men wait until the sun is setting, then build a fire on the ground below where the nest is hanging. Using a hooked stick to detach the nest from its connection site, the nest is detached from the plant and immediately placed in the fire. Once in the fire, the adult insects evacuate as quickly as possible while the larvae die as they are toasted. The combs with the toasted larvae inside are then removed from the fire and brought home. The larvae can be considered celebratory food, and when a boy is newly born it is the father’s duty to collect a nest alone.


Cultural significance of wasp stings

In several tribes from the Amazon basin, ''P. pacificus'' along with other ''Polistes'' wasps were collected so that their stings could be used as instruments of pain during cultural rights of passage. For example, the Oyana people in Surinam constructed breastplates that were made of woven plant stems, in which live wasps were trapped. During initiation ceremonies, young boys would have to wear the breast plate all night, enduring the pain of continual wasp stings. This would have been especially painful because wasps are able to retract their stings and repeatedly insert them with new venom multiple times.


References

{{Taxonbar, from=Q14423456
pacificus Pacificus was a disciple of St. Francis of Assisi, born probably near Ascoli, Italy, in the second half of the twelfth century; died, it is thought, at Lens, France, around 1234. Poet Laureate Local authors identify him with a certain William of ...
Insects of South America Insects described in 1804