Angiopolybia pallens
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''Angiopolybia pallens'' is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of social wasp predominantly found in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. The
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 ...
is generally seen in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
ian
rainforests Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainforest ...
. This species was discovered by Lepeletier in 1836. It typically feeds on
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
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
. In fact much of its feeding behavior and impact on humans is centered on feeding on animal carcasses. The wasp species displays a caste differentiation that can be seen by difference in ovarian development. Additionally they have a unique colony establishment procedure. It begins with a few individuals from the nest leaving to find a good site and then the rest of the colony follows using specific communication signals that are further discussed in this article.


Taxonomy and phylogeny

The
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
social wasp
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Angiopolybia'' contains four different species. The wasp is within the tribe
Epiponini The Epiponini (formerly known as Polybiini) are a large and diverse tribe of social wasps inhabiting the Neotropical region, with some species' ranges extending into the Nearctic region. Selected species

* ''Apoica pallens'' * ''Leipomeles d ...
and is a basal swarm-founding wasp. A swarm founding wasp is a wasp that finds its nest by sending out a group of wasps going out to find a new nest site. It is considered a
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
wasp. Most social wasps have the unique characteristics of a caste system in which the queen wasp is the primary producer of offspring. This characteristic of a caste system is evident in the behavior of ''A. pallens. ''Lepeletier originally discovered the species in 1836. He classified the wasps under a different genus ''Rhopalidia'' and when he discovered the wasps he also found another species under the name of ''R. rufithorax''. The species was later transferred to the genus ''Angiopolybia'' that shared with ''A. paranesis, A. obidensis, and A. zischkai''. These species are widespread in the
Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology c ...
and extend from
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
to
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
.


Description and identification

The ''A. pallens'' are distinguished by the
ombre Ombre (, pronounced "omber") or l'Hombre is a fast-moving seventeenth-century trick-taking card game for three players and "the most successful card game ever invented." Its history began in Spain around the end of the 16th century as a four-pe ...
-like progression from yellow to black on their bodies. They have four legs that progress from a lighter yellow to a dark orange. The wasp's body is skinny with the back coming to a point. Their wings are slim and black. The slim wings cause the ''A. pallens'' to fly low and for short distances. This means that their foraging area is limited to about 1,800 square meters. The workers, queens and intermediates of the colony are almost indistinguishable with the queens only occasionally being larger in size. The nest of ''A. pallens'' is generally in a
funnel A funnel is a tube or pipe that is wide at the top and narrow at the bottom, used for guiding liquid or powder into a small opening. Funnels are usually made of stainless steel, aluminium, glass, or plastic. The material used in its construct ...
shape with a downward facing entrance. The nest is usually attached to a
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
by a central
pedicle 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 ...
. In general the nest is strategically placed at a diagonal in order to provide more coverage from a leaf. The nest is usually made from a mixture of salivary secretions and vegetal fibers. All social wasps use plant
fiber Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...
as the primary material in the construction of their nests.


Distribution and habitat

The ''Angiopolybia pallens'' is predominantly found in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. They are among the most common species of wasps in this area by an overwhelming amount, as seen in multiple feeding behavior studies. The wasps prefer more humid environments and are more active when humidity levels are higher. However, the wasps are generally more active during the cooler parts of the day, like the early morning or around dusk. The nests of these wasps are typically found in rainforest environments in clearings or on the edge of the forest. ''A. pallens'' typically have nests that are large and easy to see hanging off of the branch of a tree. This means that their nests are subject to predation by ants and other animals. In order to compensate for the size and easy sighting the nests are protected from any ant predation by
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
ants with which they share the tree.


Colony cycle

The colony cycle begins with the pre-emergence phase where the queen and a few adult workers establish a new nest. The queen already has been fertilized and is able to produce a few workers in the beginning in order to best assist with the building of the nest. The swarm starting the nest is mostly made up of adults that are already mature before the next nesting cycle begins. Then once the nesting cycle begins, the
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 ...
and
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
are laid. Next is the worker production phase where they have the majority of eggs, larvae and
pupae A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
. This phase introduces the highest amount of reproduction and the intermediates are produced during this time. Once the nest and colony has reached maturity, the male production phase begins and the males eventually fertilize an intermediate. This fertilization distinguishes the intermediates from the queens and the colony cycle is repeated. The wasps are most active from months of July–December.


Foraging behavior

''A. pallens'' has exhibited a unique feeding pattern compared to most social wasps in a variety of ways. They are more active in the daytime compared to many other wasp species. These wasps tend to be most active at collecting nectar between 7 and 8 am, collecting prey between 10 and 11 am and collecting pulp for nest building generally happens before foraging. In general, other wasp species were found to start their activity later in the day and end earlier. ''A. pallens'' are usually the initial wasps present at the site of an animal carcass trap set by Silveira in an experiment conducted to study the wasps feeding patterns. The wasps also tend to collect nectar, prey and pulp in almost equal percentages. Other social wasps tend to focus on the collection of glucidic kinds of foods. This behavior still perplexes scientists.


Carrion Feeding

''A. pallens'' have shown high prevalence of
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
feeding in Brazil. When compared to six other species of wasps, they made up 43.5% of the wasps that approached a carcass. The wasps do not transport pieces of flesh to the nest but instead consume at the feeding site, and the eaten food is subsequently regurgitated to other workers in the nest. The wasps also are more likely to consume carrion from July to December, which are the drier months of the year. It has been claimed that this wasp is the most important carrion consuming wasp in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. An additional behavior that ''A. pallens'' exhibit is the interaction with other wasps and ants when feeding on carcasses. ''A. pallens'' are typically the first wasps to rush an animal carcass but are typically displaced when another bigger wasp species approaches the carcass. However, the wasps will hover over ants occasionally darting downwards toward them causing the ants to flee. The wasps will then land on that particular area of the carcass and feed. They continue to intimidate the ants preventing the encroachment on their space by raising their wings and darting towards the ants. In this way the wasps are successful at obtaining food from carcasses even in the presence of ants.


Swarming behavior

Swarming Swarm behaviour, or swarming, is a collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or migrating in some direction. ...
is often a method used by wasps to travel from one nest site to find a new nest site. This type of behavior is shown by ''A. pallens'' and is significant to note as it displays the importance of communication within the wasp
colony In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the ''metropole, metropolit ...
. The wasps will divide themselves into two different swarms when beginning the process of finding a new nest site. The first swarm consists of queens and workers and this group leaves the nest site to find a new site to establish a colony. They leave the site because of “cyclic intracolony factors”, meaning it is time to move on and establish a new colony. The second swarm includes the rest of the adult population and happens upon the destruction of the previous nest. It is important to note that the queen is the founder of a colony as she is needed to produce new workers and intermediates to help fully establish the nest.


Communication

Once they have found a new nest site, the ''A. pallens'' convey this information to other workers and intermediates through glandular secretions from their antennae. They make upward runs of several centimeters long on vertical surfaces providing a road map of sorts for the adult wasps to follow. The front body of the wasp will angle slightly upward while the antennae works its way upwardly to maintain the connection with the surface. The wings perk up as they make this connection. Typically the ''A. pallens'' begin to use this trail about 3 hours after the swarm formation. This means that generally the new site is already being developed. In fact the dragging behavior seen to deposit the
glandular In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
secretion 440px Secretion is the movement of material from one point to another, such as a secreted chemical substance from a cell or gland. In contrast, excretion is the removal of certain substances or waste products from a cell or organism. The classical ...
by the wasp is seen only after the deposition of building material at the new nest site.


Ants and Nest Selection

The two primary causes of nest mortality for ''A. pallens'' and other social wasps is predation and inclement weather. Therefore, the wasps attempt to find a location that will both protect them from bad weather as well as bar any access for predators.
Army ants The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limit ...
present one of the largest threats as predators for ''A. pallens'' nest sites and are considered to be one of the driving forces in the evolution of their nest architecture. ''A. pallens'' have been found to build their nests near arboreal ants in order to help protect against the army ants. However, this does not mean that the wasps avoid predation all together. An evolutionary response to this chance of attack is the ability of the adult population to escape, move the colony, and replace the nest.


Queen intermediate conflict

The wasps in ''A. pallens'' colonies can be broken up into three groups: queens, intermediates, and workers. These groups are distinguished by whether they are mated or
virgin Virginity is the state of a person who has never engaged in sexual intercourse. The term ''virgin'' originally only referred to sexually inexperienced women, but has evolved to encompass a range of definitions, as found in traditional, modern ...
, and whether they have developed or undeveloped
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
. As discussed earlier on, there is little variance in the size of the wasps depending on their caste placement. This implies that the ''A. pallens'' establishes the queen wasps during adulthood rather than during development. In other words, the post conception determination is by the conflict between individuals in the colony and not by their feeding during larval development. This means that there is a high amount of competition always present during colony formation. Due to this determination, intermediates are in conflict with the queens. The intermediates possess the ability to reproduce because they have developed ovaries. Additionally the queen invests equally in each offspring, as it is unknown whether the larvae will be deemed an intermediate or queen until later on in adulthood. This means that the intermediates have the chance of producing offspring in the next colony along with other queens, and the queens have little to no control over this reproduction.


Morphological Appearance

In females who are workers and intermediates the ovaries are all in the same stage, making them almost morphologically identical. The queen's ovaries can be distinguished by the presence of post mating
oocytes An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ge ...
. Additionally, in most colonies workers and intermediates tend to be of equal size while the queen is larger. This size difference is almost impossible to distinguish, meaning that the caste system for these wasps cannot be detected by differences in size alone. This lack of size difference implies that the queens for a colony are not determined during development. The intermediates in the colony have little to no morphological differentiation compared to the queens, making it easy for them to reproduce during the next colony cycle without regulation from the queen.


Genetic diversity

There has been significant genetic differentiation found in ''A. pallens'' populations across the
Atlantic 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 an ...
and Amazonian rain forests. Much of this differentiation has been attributed to the fact that these rain forests are separated by the country of Brazil whose border they cross. Due to this isolation, gene flow among the populations is often unlikely to occur. The ''A. pallens'' species has been separated into eight distinct
haplogroups A haplotype is a group of alleles in an organism that are inherited together from a single parent, and a haplogroup ( haploid from the el, ἁπλοῦς, ''haploûs'', "onefold, simple" and en, group) is a group of similar haplotypes that shar ...
which are further separated into groups A, B and C. The wasps in group A are from southern
Bahia Bahia ( , , ; meaning "bay") is one of the 26 Federative units of Brazil, states of Brazil, located in the Northeast Region, Brazil, Northeast Region of the country. It is the fourth-largest Brazilian state by population (after São Paulo (sta ...
on the eastern side of Brazil with group B combining haplogroups found in northeastern Bahia,
Santa Terezinha Santa Teresinha is a municipality in the state of Bahia in the North-East region of Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin Ameri ...
, and Vera Cruz (also all on to the east of Brazil.) Group C included wasps found in
French Guiana French Guiana ( or ; french: link=no, Guyane ; gcr, label=French Guianese Creole, Lagwiyann ) is an overseas departments and regions of France, overseas department/region and single territorial collectivity of France on the northern Atlantic ...
,
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
and
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
. The presence of these different haplogroups suggests that at one point the species had a large range of land in which it could inhabit. The hypothesis that forests were connected before the glacial events is supported by the presence of ''A. pallens'' in such widespread regions of South America. The species likely began to diversify due to geographic isolation from
Savannas A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
in the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
era.


References

{{Taxonbar, from=Q14423278 Vespidae Hymenoptera of South America Insects described in 1836 Taxa named by Amédée Louis Michel le Peletier