Vespula infernalis
Vespula infernalis is an obligate parasitic wasp, parasitizing the nests of other species in the genus "Vespula". Its common host species is "V. acadica" in North America. It is sometimes called the cuckoo yellowjacket wasp due to its inquiline lifestyle. They differ from other parasitic wasps in their intensely aggressive behaviour during invasion and occupation of the host colony. Several morphological adaptations such as bigger body parts and more curved sting shafts are observed in these wasps to aid their aggressive parasitic behaviour. Once they occupy a host's nest, "V. infernalis" are known to engage in mauling  and chasing of host workers and forced trophallaxis. Female wasps will also force host workers to feed and take care of their brood . Taxonomy and phylogeny. "Vespula infernalis" is a member of the family Vespidae. Furthermore, "Vespula infernalis" is nearly identical in appearance and biology to the Old World species "V. austriaca" and is considered its sister species. Until recently the Nearctic population was considered to be the same species, and appears in all of the literature before 2012 as "V. austriaca", though some sources are not yet updated (e.g.). Description and identification. This wasp is an obligate social parasite . As a result, it possess special morphological adaptations to take over host colonies. These adaptations include larger body parts such as a larger head width than its host "Vespula acadica". Because of its parasitic lifestyle, "V. infernalis" has a large stinger with very curved sting shafts; this curving allows the parasitic wasp to puncture through the intersegmental membranes of host colony workers who are defending their nests. The male abdomen is longer with thick lateral black bands while the female abdomen is wider with fewer lateral black bands and black dots. "V. infernalis" wasps have a characteristic black and yellow  colouring throughout their bodies. Irregular margins and narrow yellow marks on the scape are often used to identify "Vespula infernalis". Distribution and habitat. "Vespula infernalis" is found in the Nearctic region, in the Boreal region of North America. The species is not considered rare and is stable throughout its range. These wasps are seen throughout all the provinces and territories of Canada and in certain states in the United States; Alaska, New Jersey, Michigan, Wisconsin, North Dakota, New Mexico, Arizona, and California. Colony cycle. This wasp is known as a social parasite. Their life cycle depends on invading and usurping  other wasps' colonies. As an obligate parasite, "Vespula infernalis" lacks a worker caste and cannot build their own nests or feed their own offspring. "V. infernalis" females will hibernate longer than their host queens, ensuring that the hosts' nests will be ready when they leave their hibernation spots. After killing the host queen, the female "V. infernalis" tends to be aggressive towards the host workers who rear her brood. These wasps have very short seasons with flight periods from June to mid July and August to mid September. Parasitic behaviour. "Vespula infernalis" parasitises "Vespula acadica". Near the dorsal edge  of "V. infernalis" mandibles, there is a fourth marginal tooth whereas the host, "V. acadica" has three. When researchers dissected individuals of "V. infernalis", they found that they had larger, sturdier mature muscle bundles in abdominal sterna and tergum than the muscles in a queen of "V. acadica". As a result, it was more difficult to dissect "V. infernalis". Nests of "V. acadica" were found to be smaller than normal due to the parasitism of "V. infernalis". Invasion. Queen-reared worker stage. When a female "V. infernalis" invades a host's nest such as "Vespula acadica "during the queen-reared  worker stage, the usurpation can be divided into two phases. The first phase is preusurpation. During this phase, interactions between the intruder and the host queen could be relatively calm and civil or aggressive. The queen may ignore the intruder and refrain from firing an attack first. Likewise, the workers exhibit a similar behaviour like their queen. But if the intruder attacks first by mauling or attempting to sting a worker, interactions between the queen and intruder become intense. The parasite, if successful, will use its stinger to paralyze the head, thorax, and legs of the host queen. This stage  is considered the most vulnerable and hence, the most popular time to invade. It is susceptible to invasion because both the queen and workers are less defensive. Post-emergence colony stage. During this stage, the colony is larger. In one experiment, a "Vespula infernalis" female was allowed to invade a "Vespula atropilosa" colony. Different from the queen reared worker stage, the workers in this stage were much more defensive; more aggression was observed in the form of stinging, mauling and chasing. Additionally, the parasite displays more aggression than the host's queen. During a fight between the parasite and the  queen, the parasite will do anything to harm or gain a better position by using her mandibles and stinger. Injuries to the queen include but are not limited to broken antennae and injured fore and hind-leg. Surprisingly, the host queen refrained from using her mandibles whereas the workers were quick to use theirs in defense. Post-invasion: parasite and host worker interactions. "Vespula infernalis" are very hostile and aggressive in invading and killing or driving away the host queen. Once the invasion of a host nest is successful, "V. infernalis" will take over the nest, inspecting and patrolling the  cells. The host workers will initially avoid the parasite and often ignore any colony duties. In response, the parasite will maul and chase the workers. While there is still tension, the inquiline typically does not sting the workers and does not engage in egg eating (oophagy). A suggested strategy may be to kill older workers of the host colony first and leave the younger workers behind to take care of the brood. During this period, a female will assert her dominance over the younger workers. This obvious display of dominance by the parasite over the host workers is more common  during early colony occupations and absent later on. The parasite displays mauling behaviour: it grabs the workers with its legs and chews on their dorsums. Furthermore, the parasite also displays mauling behaviour after trophallaxis, which it initiates by soliciting and chasing a worker. "Vespula infernalis" rarely forage, build cells, or feed larvae, nor do they help thermoregulate (workers fan during hot temperatures or adopt warming postures during low temperatures). Furthermore, even when a nest is attacked by enemies, the parasite will remain inside the nest rather than help the workers defend it. Suppression of reproduction of host workers. Several signs  indicate that "Vespula infernalis" repress the reproduction of host workers. There is no worker oviposition in nests of "Vespula acadica" parasitised by "Vespula infernalis". Secondly, if workers were ovipositing then there would be higher levels of aggression between workers. However mauling was reduced between workers. The frequent aggressive and assertive behaviours of the parasites aid in asserting reproductive monopoly. Another indication is that parasites are observed to patrol cells frequently, hence defending oviposition sites. Lastly, during gastral dragging (dragging of abdomen across comb), substances that prevent oogenesis in workers are released from the Dufour's gland of the parasite. Diet . While other wasp queens depend on secretions from larvae for food, "Vespula infernalis" obtain their nourishment in the liquid form through solicitations and trophallaxis of host workers. They rarely solicit larvae for nourishment. The parasites will often use force to obtain what they need. During forced feeding, the parasite will grab the host worker using its forelegs to reel in the worker. More forceful behaviour includes pushing the host worker against a comb of the nest. While some parasitic behaviour (i.e. maulings) only occurs during early post-invasion stages, forced trophallaxis behaviour occurs throughout. Towards the later stages, the intensity  of forced feeding declined, leading to host workers escaping forced encounters. Usually forced trophallaxis encounters were short lasting from 1–5 seconds. But some encounters lasted for as long as 43 seconds. Characteristics of the venom. Due to the parasitic lifestyle of "Vespula infernalis", the sting is a vital apparatus. It consists of the shaft, aculeus, Dufour's gland, and the venom glands. The Dufour's gland and venom reservoirs are the biggest among "Vespula infernalis" females. In an experiment where the oily venom extracted from "V. infernalis" was injected into "Vespula pensylvanica" workers, the lethality (LD50) value was between 20  and 30 mg/kg. "V. pensylvanica" queens had a higher tolerance to "V. infernalis" venom than workers (LD50=81 mg/kg). On average, it took 18.6 μg of venom to kill about half of the queens and 2–3 μg of venom to kill half the workers. The venom has the ability to paralyze and impair activities such as flying. Material obtained from the Dufour's gland was also injected into " V. pensylvanica" wasps; no significant effects were observed, indicating the gland does not contain a poison. Generally, in order to kill a host worker, a "V. infernalis" has to use about  a third of its venom supply. More venom is needed to kill a queen. Surprisingly, "V. infernalis" venom is as lethal as the venom from "V. atropilosa" and "V. pensylvanica". Reproduction. In preusurpation "V. infernalis" individuals, Dufour's gland contains clear oily material. It is part of the reproductive organs along with the ovaries and a poison gland. Distinctly enough, in new fall wasps, the Dufour's gland was empty and flat and in older wasps, the gland had a little oil but was flat as well. Hypertrophy of the Dufour's gland is known to occur but more evidence  is needed. During and after the mauling period, female "Vespula infernalis" will oviposit. This time period is brief. The females insert eggs in new cell combs. Workers act no differently during period of oviposition. However, one instance of a worker interrupting the oviposition of the parasite was recorded, which led to mauling of the worker by the parasite and the egg falling out of the designated cell. Eggs laid by "Vespula infernalis" are not only found in cells but also found on the cell wall, exterior of the comb, between cells, or along the caps.