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''Dolichovespula adulterina'' is a species of
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
social wasp found in the Palearctic region. ''D. adulterina'' feeds on a variety of foods, including insects, spiders,
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 ...
, meat,
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is estim ...
, fruit, nectar, and larval secretions. ''D. adulterina'' was formerly considered to be synonymous with ''D. arctica'' from the Holarctic region, but more recent research indicates that ''D. arctica'' is a separate species.


Taxonomy and phylogeny

''D. adulterina'' is a parasitic wasp in the order Hymenoptera. It is a member of the subfamily Vespinae and was labeled a subgenus of the genus ''
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 ...
'' at one point. ''
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' ...
'' spp. are identified by a gently notched labrum on the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
border. This is the location of sensory bristles. ''Dolichovespula'' spp. exhibit microscopic spinous growths all over the palate region, whereas other species have localised growths. Another distinctive feature is the monodentate mandible, where the primary incisor is always linear. Its body size is nearly identical to species of'' Vespula''.


Identification

The length of the forewing is 13.0–14.0 mm in females and 10.0–12.0 mm in males. Commonly, its body is an ivory colour, but it can be pale yellow on occasion; the pale yellow subspecies are found in the western Nearctic and Palearctic regions. Usually, the clypeus is elongated, but sometimes it will be small and round. Females almost always have a black, disc-like spot, whereas males have a black mark that extends to the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
margin of the clypeus. Melanic individuals have a black spot extending to the dorsal margin. The postocular band in females is narrower relative to the
lateral Lateral is a geometric term of location which may refer to: Healthcare *Lateral (anatomy), an anatomical direction *Lateral cricoarytenoid muscle *Lateral release (surgery), a surgical procedure on the side of a kneecap Phonetics *Lateral cons ...
postocular stripes, whereas in males it is rarely narrowed. Occasionally, small ivory sports are located on the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum (ventral), and the propleuron ( lateral) on ...
behind the pronotal fovea. Females present with pairs of black discal spots on
tergum A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'ma ...
5, and occasionally on
terga A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
3 and 4, as well. In males,
tergum A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'ma ...
6 usually is black with a less developed fascia than previous terga. Furthermore, in males,
tergum A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'ma ...
7 is usually black and rarely contains two ivory spots. A black and white, sometimes pale yellow, this wasp has brown-tinted wings. Normally, abdominal segments 1-5 are bordered, towards their rears, with a thin, white band. The large, dark malar space between the eye and the yellow jaw puts this species in ''Dolichovespula'' rather than the related genus ''
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 ...
''.


Distribution and habitat

''D. adulterina'' has a Palearctic distribution, and it parasitises '' D. saxonica'' and '' D. norwegica''. ''D. saxonica ''constructs nests in trees and shrubs no more than 2 m above the ground. Sometimes, ''D. saxonica '' nests in beehives, underneath overhanging edge of roofs (sometimes up to 7 m above ground) and porches, and cavities of walls. On occasion, they may also nest in tree holes or holes in the ground.


Biology

''D. adulterina's'' main host in Europe is ''D. saxonica'', but it also parasitises ''D. norwegica''. This species produces no workers; the eggs laid in the host's nest produce queens and males, which are nurtured by the workers of the host species.


Colony cycle

''D. adulterina'' is a parasitic species, so its lifecycle is consistent with the host it parasitises. Furthermore, ''D. adulterina'' invades the nest of host species before the first set of workers appears, and lives alongside the queen for a time before usurping her.


''D. saxonica''

''D. adulterina'' most often parasitises ''D. saxonica''. In mid-May or just before, colonies initiate. The first set of workers emerges in early June and begins working on large cells mid-June. Queens emerge in early July and the colony continues until just after mid-August. Regardless of the host species, the queen ''D. adulterina ''invades the host colony around the same time that the first workers start to emerge. The invasive queen waits around 10 days before killing the host queen and laying her eggs. Males emerge in late July and new queens emerge in early August. After a few days, the males and females leave for their mating flights. The colony then terminates in mid-August. The
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
queen dies by mid-July, if not earlier, from fights with host workers. After the colony ceases, the queen enters hibernation in a sheltered area. She will then emerge and feed on nectar and malaxate
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 ...
until she attempts to usurp a host colony. On average, the colony cycle for a successful parasitic queen is about 2.8 months.


Parasitic behaviour

''D. adulterina'' is a parasitic social insect. Social insects are labeled
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
when the parasitic female enters the host nest and assumes the reproductive responsibility of queen while simultaneously leading the pre-existing workers. ''D. adulterina ''is an
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
species, meaning that workers are not produced. One of its hosts species is '' D. norwegica'', although this was not accepted by some authorities for some time, indirect evidence was the presence of ''D. adulterina'' in regions where ''D. saxonica'' was absent but the related ''D. norwegica'' was present and this was confirmed by direct observation. Furthermore, they do not have a worker caste and they do not construct nests. Instead, they will use the host nests and workers to raise their offspring. ''D. adulterina '' uses a submissive approach while the host nest is under attack, living alongside the host queen for a while before attempting to kill her. This is different from other inquiline species, such as '' V. austriaca'', which attack the nest's queen immediately. During its time in the nest, ''D. adulterina'' is an active member of the colony, feeding larvae and constructing portions of the nest. Workers of the host species were observed interacting with ''D. adulterina'', suggesting that the parasite may emit a chemical that pacifies the host. This is also supported by the presence of an enlarged
Dufour’s gland Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed at ab ...
relative to other vespine species. Time of invasion also differs between species. ''D. adulterina'' invades the nest of the host species before the first set of workers appears, whereas '' Vespula austriaca'' invades after the emergence of the first set of workers.


Diet

''D. adulterina'', like other social wasps, is primarily
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
. The species feeds on a multitude of insects, spiders, and arthropods. To do this, adults malaxate prey and feed on the juices. Other times, they have been observed eating meat, molluscs, fruit, and nectar. Larvae produce a secretion on which adults feed, as well. Wasps limit themselves to attacking live prey, but other species have been observed to feed on carcasses.
Trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth ( stomodeal) or anus-to-mouth ( proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pher ...
is a common process among wasps, where one member transfers food to another member by feeding it mouth-to-mouth. Members of the host nest are mainly responsible for feeding the parasitic ''D. adulterina ''larvae.


Sting

''D. adulterina's'' stinger is an adaptation to its parasitic biology. The
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
of the stinger is thicker than average and is curved. These modifications allow for ''D. adulterina ''to effectively penetrate the intersegmental membranes of the host. ''D. adulterina '' is better equipped defensively, as well. The abdominal
sclerites A sclerite (Greek , ', meaning " hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly ...
are closer together, providing a smaller gap for incoming stingers to penetrate, thereby decreasing chances of injury.


Emery's rule

''D. adulterina'', along with ''V. austriaca'', ''D. arctica'', and ''D. omissa'', are the only four vespine wasp species that display inquilinism. Furthermore, all of these species abide by
Emery's rule In 1909, the entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.Emery, C. "Über den Ursprung der dulotischen, parasitischen und my ...
, which states that a parasite is more closely related to its host than to any other species. In host colonies, ''D. adulterina ''queens have allomonal control over worker wasps. While ''D. adulterina ''is able to comfortably coexist with its host, it does not integrate nearly as well into host colonies as inquiline ant species. Due to its mediocre proficiency, inquiline wasps are only able to parasitize other species that exist in small colonies and have short cycles.


References

{{Taxonbar, from=Q916125 Vespidae Hymenoptera of Europe Taxa named by Robert du Buysson Insects described in 1905