Polistes semenowi

Polistes semenowi is a kleptoparasitic paper wasp that is found in several regions of high altitude in Europe. As one of only three obligate parasites in the genus Polistes, it uses the nests of other paper wasps (primarily Polistes dominula) to rear its young.^[1] In order to evade detection by the host nest, P. semenowi employs mimicry by adjusting its cuticular hydrocarbons to match those of the host.^[2] Once the host nest has been infiltrated, the parasitic female will physically attack the host queen in order to subdue her and become the colony's new queen. P. semenowi displays several morphological adaptations for parasitism such as increased mandible size and an enlarged Van der Vecht's organ. This species is unusual because it does not have the ability to produce workers and is only able to produce individuals who have the capacity to reproduce..

Taxonomy and phylogeny

Due to its morphological differences from other Polistes species, Polistes semenowi was placed in a separate genus Sulcopolistes by Blüthgen in 1938.^[3] However, in 1991 Carpenter established that this species belonged in the genus Polistes.^[1] P. semenowi is closely related to Polistes sulcifer and Polistes atrimandibularis, which are the only other obligate social parasites in the genus Polistes.^[4] Research using mitochondrial rRNA suggests that these three species descended from a common ancestor, and that they are more closely related with Polistes nimpha and Polistes dominula than with Polistes gallicus and Polistes biglumis.^[1]

Description and identification

P. semenowi is larger than most Polistes species, which initially caused it to be classified in a separate genus (Sulcopolistes).^[3] Both the first femur and posterior tibia of this wasp are elongated, and its mandibles are significantly thicker than those of other wasp species. Moreover, the mandibles of this species, similarly to Polistes sulcifer and Polistes atrimandibularis, are marked by a distinct groove.^[1] This wasp species also has distinctive black markings on its clypeus, the function of which is currently unclear.^[5]

Distribution and habitat

P. semenowi populations typically exist near regions of high altitude around the Mediterranean basin. However, on occasion, they may also be found around the Caspian basins. The distribution of P. semenowi is patchy as a result of its altitudinal migration patterns. During the winter, the wasps ascend to a higher altitude; in the spring, however, they descend to the lowlands to find host colonies.^[1]

Colony cycle

These wasps migrate to high altitudes to mate and then proceed to overwinter in the same mountainous areas.^[2] In the spring, females move down the elevation gradient in order to parasitize P. dominula, a lowland species. Once a host nest has been discovered, the P. semenowi female will attempt to usurp the nest. The timing of this usurpation is intimately linked to the emergence of P. dominula workers. If no workers have emerged, the hosts may simply abandon the colony; if most of the workers have already emerged, they may be able to fend off a P. semenowi invasion.^[1] Having usurped the dominant female host, the P. semenowi female proceeds to lay her eggs. After a period of several weeks, the parasitic female will abandon the nest.^[3] Once her offspring emerge from the host nest, they will migrate to high altitudes, continuing the cycle.

Parasitism

P. semenowi are one of only three species of Polistes that are obligate social parasites (along with P. sulcifer, and P. atrimandibularis). This means these wasps take advantage of the social systems of other species.^[4] As a social parasite, P. semenowi usurps a host colony in order to take advantage of the entire host colonial cycle.^[1] Specifically, this parasitic wasp exploits P. dominula, a lowland wasp species. Although P. semenowi is a specialized parasite of P. dominula, it is also able to parasitize colonies of P. nymphus. Since P. semenowi is unable to create nests and produce worker classes, it is completely dependent on a host colony to fulfill these functions.^[4]

Because P. semenowi normally attacks the host's nests just prior to worker emergence, it is surmised that such selection pressure might have impacted the developmental times in P. dominula to be shorter. It would be advantageous for the hosts to make their brood developmental time to be shorter so that they can rapidly generate workers and defend their nests.

Evolutionary basis of parasitism

Like other Hymenopterans, Polistes species undergo a complete metamorphosis during development in which the young are completely dependent on workers for all of their food and protection needs. A large energetic investment is necessary in order for the young to successfully pupate and grow to reproductive age. The amount of resources invested in the young determines which caste they will belong to: less food will cause the larvae to develop into workers, while more food will cause the larvae to develop into reproductives.^[1] By manipulating another species into investing a large amount of energetic resources into their own young, P. semenowi would be able to take advantage of host workers and would no longer have a need to produce its own workers. Instead, more resources could be devoted to larvae, causing all of them to develop into reproductives. Over time, P. semenowi has completely lost the ability to create its own worker class and has adopted a strategy of parasitism.^[4][2] The parasitic larvae display rapid growth, which allows P. semenowi to optimize its rate of offspring production.^[1]

Establishing dominance over host queen

Approximately two months after a P. dominula female has founded a colony, the female parasite will attempt to usurp the nest.^[2] Approaching the dominant host female, the parasite will aggressively attack her in an attempt to drive her out of the nest.^[6] Behaviors include chasing the queen out of the nest as well as physically confronting her.^[3] These behaviors have been demonstrated in the laboratory: in controlled experiments, female parasites introduced to host nests immediately entered the nest and identified the most dominant female, proceeding to aggressively approach her.^[2] After initially physically establishing dominance over the queen, the parasitic female will coexist with subordinate females and may in some cases allow the former dominant female to stay in the nest.^[1]

Reproduction within host colony

Once she has entered the colony and subdued the dominant host female, the parasite immediately begins to start laying eggs. Some time after the initial invasion – approximately thirty days later – the hosts will chemically recognize the parasites as their own species.^[2] Additionally, the parasite will attempt to destroy the larvae and eggs of the host queen in order to ensure that the host colony invests all of its resources into her young. After this initial destruction, however, the P. semenowi female will not attack any emerging host workers and will spend most of her time laying eggs.^[3][1] Unable to differentiate between the parasite and their own species, host workers will feed and care for the parasite young as the parasite queen continues to lay eggs.

Morphological adaptations for parasitism

In order for their parasitic strategy to be successful, P. semenowi must be able to enter and successfully usurp a host nest. However, host colonies are able to mount a defense by mobilizing any emerging workers as well as dominant females. Therefore, P. semenowi must be morphologically adapted to overcome such defensive attacks. The parasitic wasp is significantly larger than its host; in fact, P. semenowi is so much larger than P. dominula that it was initially thought to be in a separate genus.^[3] The first femur and posterior tibia of this wasp is enlarged, providing an advantage during attacks. P. semenowi also has thicker and larger mandibles, which are used during attacks to drive off dominant host females. However, since this species does not use its sting during a host colony invasion, its sting is not morphologically distinct from other Polistes species.^[1]

Mimicry

Host colonies will only mount a defense against parasites if they recognize it as distinct from their own species. Although the parasitic wasp is able to overcome the physical defenses of a host nest, it must also be able to camouflage itself after usurpation. This aids the wasp from having to continually battle for dominance. For P. semenowi, this is achieved by mimicking the chemical signals of the host colony so that the parasite is recognized as a member of the host species. By successfully mimicking the hydrocarbon patterns of host wasps, P. semenowi wasps are accepted by the colony and can ensure that host workers will raise the parasitic offspring.^[4]

Hydrocarbon signals

Like other social insects, wasps recognize each other through chemical signals. Each colony has a specific blend of hydrocarbons that is secreted onto the cuticle of the wasps.^[2][7] This hydrocarbon signature allows individuals to distinguish nest mates from interlopers. An unfamiliar hydrocarbon signature signals that the colony should mount a defense against a potential intruder. As soon as a female P. semenowi wasp enters a host nest, she begins to vigorously rub her abdomen against the comb.^[1] This allows the parasitic female to coat herself with the host hydrocarbon pattern, which she immediately begins to mimic. Although P. semenowi's initial hydrocarbon signature is relatively close to that of its host P. dominula, after usurpation, the species is able to exactly match the hydrocarbons of its host colony.^[2]

Van der Vecht's organ

The hydrocarbons secreted by Polistes wasps are efficiently spread on to the cuticle by a structure called the Van der Vecht’s organ. This structure is located on the last gastral sternite of the wasp, on the anterior edge. It is composed of a hairy, transparent cuticle. When compared to females of their host species P. dominula, female P. semenowi wasps have a significantly enlarged Van der Vecht’s organ. Since the Van der Vecht’s organ is used to spread hydrocarbons which enable P. semenowi mimicry, the increased size of the structure in this species most likely occurred as a result of selective pressure on the wasp due to the success of its parasitic life strategy.^[4]

References