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How do *Polistes sulcifer* (cuckoo paper wasp) eggs hatch so quickly?

How do *Polistes sulcifer* (cuckoo paper wasp) eggs hatch so quickly?


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The Polistes sulcifer wasp is a brood parasitic species, which parasitizes colonies of the paper wasp Polistes dominulus. During an extremely brief time window in the spring, just before the P. dominulus workers have emerged, the P. sulcifer female will invade a P. dominulus nest. Once inside, it will kill or evict the host queen, ensure its cuticle hydrocarbon profile will not lead to its being attacked, and then lay its own eggs. It will rely on the host workers to tend to these.

(Note: I don't know whether it just alters its own hydrocarbon profile, or whether it also alters that of the nest. It's tangential to this question.)

The P. sulcifer wasp needs to time its attacks precisely. If it invades too late, the workers will have emerged and the colony will have a lot more wasps to defend it. If it invades too soon, there will not be enough host workers emerging to tend to its brood, especially if the host queen has not laid many eggs. P. sulcifer itself does not have a worker caste, and so is entirely reliant on host workers.

Research published in 2004 (paywalled, unfortunately) showed that the parasite wasp eggs took an average of 5 days to hatch after being laid, with the minimum observed being 3 days. This was in marked contrast to the host eggs, with average time 8.6 days in an unparasitized host colony, and 9.1 in one which was parasitized. Likewise, the parasite larval stage was much shorter, lasting an average of 9.8 days as opposed to 16.6 for host larvae in a parasitized nest (14.8 in an unparasitized colony) and the pupal stage also slightly shorter.

The parasite larvae were able to attract more attention from the workers than the host larvae - including being fed more often - which would appear to offer an explanation for the faster larval development and shorter larval/pupal stages. But this does not explain the shorter egg stage. Rapid development is clearly advantageous in terms of maximising the number of offspring produced, so a shorter egg stage is advantageous to the parasite, but the mechanisms behind this short incubation period are not known.

In the famous example of an avian brood parasite, the cuckoo, the parasite eggs also hatch more rapidly than the host eggs. This is, at least in part, because the female cuckoo holds the egg in her oviduct for an additional 24 hours prior to laying, at a higher temperature than an egg being incubated in the host nest. This gives the cuckoo egg the equivalent of a 30 hour head start on the host eggs.

In the aforementioned 2004 research paper, it was speculated that the wasp might use a similar mechanism, but this was not known, and no research into this had been carried out. A later 2006 paper also stated that no information was known about this, and I have been unable to find any further research on the topic.

Has there been any subsequent research into how P. sulcifer has been able to ensure its eggs hatch abnormally soon after being laid?

Sources - paywalled:

Cervo, R., Macinai, V., Dechigi, F., & Turillazzi, S. (2004). Fast growth of immature brood in a social parasite wasp: a convergent evolution between avian and insect cuckoos. The American Naturalist, 164(6), 814-820.

Ortolani, I., Turillazzi, S., & Cervo, R. (2008). Spring usurpation restlessness: a wasp social parasite adapts its seasonal activity to the host cycle. Ethology, 114(8), 782-788.

Sources - non-paywalled:

Cervo, R. (2006, January). Polistes wasps and their social parasites: an overview. In Annales Zoologici Fennici (pp. 531-549). Finnish Zoological and Botanical Publishing Board.

Cervo, R., Dani, F. R., Cotoneschi, C., Scala, C., Lotti, I., Strassmann, J. E.,… & Turillazzi, S. (2008). Why are larvae of the social parasite wasp Polistes sulcifer not removed from the host nest?. Behavioral Ecology and Sociobiology, 62(8), 1319-1331.

Other sources: Davies, Nick B. (2015). Cuckoo: Cheating by Nature. Bloomsbury. ISBN 978-1-4088-5658-1.


Polistes

Wasps of the cosmopolitan genus Polistes 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 for this genus in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies. Their innate preferences for nest-building sites leads them to commonly build nests on human habitation, where they can be very unwelcome although generally not aggressive, they can be provoked into defending their nests. All species are predatory, and they may consume large numbers of caterpillars, in which respect they are generally considered beneficial. The European paper wasp, Polistes dominula, was introduced into the US about 1981 and has quickly spread throughout most of the country, in most cases replacing native species within a few years. This species is very commonly mistaken for a yellow jacket, as it is black, strongly marked with yellow, and quite different from the native North American species of Polistes. The cuckoo wasp, Polistes sulcifer, is an obligate social parasite, whose only host is P. dominula. Polistes annularis, whose species name is Latin for "ringed", is also known for its distinctive red body color. Polistes metricus adults malaxate their insect prey by chewing them into a pulp, sucking out and ingesting the body fluids, then feeding the rest of the morsel to their larvae. The most widely distributed South American wasp species, Polistes versicolor, is particularly common in the southeastern Brazilian states. This social wasp is commonly referred to as the yellow paper wasp due to the distinct yellow bands found on its thorax and abdomen. Polistes wasps can be identified by their characteristic flight their long legs dangle below their bodies, which are also more slender than a yellow jacket.

2.2. Lifecycle Worker phase
The worker phase usually begins in the early summer, roughly two months after colony initiation, with the emergence of the first workers. These new females take up most of the colonys work duties, foraging, caring for brood, and maintaining the structure of the nest. Around this time, those females which assisted in nest foundation if present are driven from the nest by aggressive behavior on the part of the foundress, and leave either to start their own late-season nests or usurp anothers.

2.3. Lifecycle Reproductive phase
The reproductive phase of the colony begins when the first female reproductives the gynes emerge from their brood cells. These reproductives differ from their worker sisters by having increased levels of fat stores and cryoprotectant carbohydrate compounds allowing them to survive the overwintering period. These reproductives contribute genes directly to the next generation, while their worker sisters normally pass along their genes indirectly.

2.4. Lifecycle Intermediate phase
Once male reproductives emerge and both males and females disperse from the natal nest for mating flights, the so-called intermediate phase begins. Brood care and foraging behavior decline and worker numbers drop as dying individuals are no longer replaced by new ones. Intracolonial aggression increases and the social cohesion of the nest declines. In temperate Polistes species, individuals almost exclusively inseminated females gather in groups of up to 50 individuals and seek a sheltered location called a hibernaculum in which to overwinter.

3.1. Behavior Kin selection
The reproductive behavior of Polistes wasps provided some of the first evidence for the mathematical biologist W. D. Hamiltons 1964 theory of kin selection. Hamilton showed that animals such as workers could be expected to provide assistance to relatives such as their queens according to the costs and benefits involved K and their degree of genetic relatedness r, and gave the rule that now carries his name, K 1/r. Early caution existed among researchers as to whether social insects could really assess their relatedness. Hamilton himself suggested an alternative possibility, namely that kin could become associated simply by "population viscosity" - that offspring tend not to disperse far from their birthplaces - and West-Eberhard 1969 found some evidence for this in Polistes. However, Polistes species are now known to learn and remember chemical signals hydrocarbons picked up from the nest to distinguish nestmates accurately from nonrelatives.


Reproductive Dominance Strategies in Insect Social Parasites

In eusocial insects, the high cost of altruistic cooperation between colony members has favoured the evolution of cheaters that exploit social services of other species. In the most extreme forms of insect social parasitism, which has evolved multiple times across most social lineages, obligately parasitic species invade the nests of social species and manipulate the workforce of their hosts to rear their own reproductive offspring. As alien species that have lost their own sociality, these social parasites still face social challenges to infiltrate and control their hosts, thus providing independent replicates for understanding the mechanisms essential to social dominance. This review compares socially parasitic insect lineages to find general trends and build a hypothetical framework for the means by which social parasites achieve reproductive dominance. It highlights how host social organization and social parasite life history traits may impact the way they achieve reproductive supremacy, including the potential role of chemical cues. The review discusses the coevolutionary dynamics between host and parasite during this process. Altogether, this review emphasizes the value of social parasites for understanding social evolution and the need for future research in this area.

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Lifecycle

The general lifecycle of Polistes can be divided into four phases: [11]

  1. Founding (or pre-emergence) phase
  2. Worker phase
  3. Reproductive phase
  4. Intermediate phase

Founding (or pre-emergence) phase

The founding stage begins in the spring when a solitary female (the "foundress") (or a small group of related females) initiates the construction of a nest. The wasps begin by fashioning a petiole, a short stalk which will connect the new nest to a substrate (often the eave of a house or outbuilding), and building a single brood cell at the end of it. Further cells are added laterally in a hexagonal pattern, each cell surrounded by six others. Although nests can achieve impressive sizes, they almost always maintain a basic shape: petiolated (stellocyttarous), single-combed, unprotected, and open (gymnodomous).

Eggs are laid by the foundress directly into the brood cells and are guarded by the foundress and the assisting females (if present). After the first larvae hatch, the foundress feeds them via progressive provisioning, bringing softened caterpillar flesh to the larvae multiple times throughout their development (as opposed to the one-time provisioning seen in some other hymenopteran groups). Each of this first seasonal brood of new paper wasps is exclusively female and destined to a subordinate worker position inside the nest they do not found their own nests and instead assist their mother in the care and maintenance of future sisters.

Some foundress wasps do not build their own nests, but rather attempt to usurp that of another female. These usurpation attempts may or may not be successful, but almost always result in impressive displays of aggression and violence. Females may also adopt a more peaceful alternative reproduction strategy by joining the nest of a close relative (usually a sister) and working as assisting females. In the latter case, such cofounding females are generally, but not exclusively, close relatives. [11]

Worker phase

The worker phase usually begins in the early summer, roughly two months after colony initiation, with the emergence of the first workers. These new females take up most of the colony's work duties, foraging, caring for brood, and maintaining the structure of the nest. Around this time, those females which assisted in nest foundation (if present) are driven from the nest by aggressive behavior on the part of the foundress, and leave either to start their own late-season nests or usurp another's.

Reproductive phase

The reproductive phase of the colony begins when the first female reproductives (the gynes) emerge from their brood cells. These reproductives differ from their worker sisters by having increased levels of fat stores and cryoprotectant carbohydrate compounds (allowing them to survive the overwintering period). These reproductives contribute genes directly to the next generation, while their worker sisters normally pass along their genes indirectly.

Intermediate phase

Once male reproductives emerge and both males and females disperse from the natal nest for mating flights, the so-called intermediate phase begins. Brood care and foraging behavior decline and worker numbers drop as dying individuals are no longer replaced by new ones. Intracolonial aggression increases and the social cohesion of the nest declines. In temperate Polistes species, individuals (almost exclusively inseminated females) gather in groups of up to 50 individuals and seek a sheltered location (called a hibernaculum) in which to overwinter.


Description and identification

P. annularis, as a member of the subgenus Polistes (Aphanilopterus), has a narrow first metasomal segment and bright orange antennal segments. Within its range of the eastern United States, these traits are shared with its allies P. exclamans and P. bahamensis . The coloration of P. annularis typically includes a ferruginous (rust-red) head and thorax and a mostly black abdomen with a single, prominent yellow ring and the end of the first segment. [1] There is geographical variation in coloration between northern and southern populations. In the north, the thorax of P. annularis has ferruginous (rust-red) markings on a predominantly black background, while in the south, the thorax is mostly ferruginous, with black markings. The legs also vary from black to ferruginous. [12] In terms of size, the fore wings are 18.5󈞃.5   mm (0.73𔂾.93   in) long in females, and 17.5󈝿.5   mm (0.69𔂾.77   in) long in males. [12]

P. annularis can be separated from darker members of P. bahamensis, a species only overlapping from Florida to Louisiana and North Carolina, by the lack of yellow markings on its mesopleuron, less developed yellow markings on the mesosoma, and the lack of additional apical yellow bands on the tergum. Similarly colored members of the subgenus Fuscopolistes, including P. metricus and darker forms of P. fuscatus , can readily be separated by the lack of contrasting orange antennal tips and the wider first metasomal segment. [12]

While many other North American Polistes species show sexual dimorphism in coloration, P. annularis and P. erythrocephalus rather uniquely do not. [12] Within these species, the males lack extensive yellow markings on the face, and instead both males and females are red-faced. Instead, sexual determination must rely entirely on structural differences. The females are identified by having 12 antennal segments and 6 abdominal segments whereas the males are identified by having 13 antennal segments and 7 abdominal segments. [12]

In the genus Polistes , the lateral mandibular groove is smaller in size than in other genera of social wasps. This groove is associated with the mandibular gland and sac-like gland reservoir used for salivary production. [16] Their larvae have labial glands, which produce silk. [17]


Distribution and habitat

P. bellicosus generally establish colonies within Texas, though the range has been observed to include North Carolina and Florida. [7] Colonies naturally occur on Baccharis sp., Ilex vomitoria , and Rubus sp. [1] in native prairies at Brazos Bend State Park, near Houston, Texas. [8] Several other paper wasp species—including Polistes exclamans , P. dorsalis , P. metricus , and P. carolina —are found in Brazos Bend State Park due to the multiple types of habitats present, including native shortgrass prairie and oak forest. [9]

North Carolina is a state located in the southeastern region of the United States. North Carolina is the 28th largest and 9th-most populous of the 50 United States. North Carolina is bordered by Virginia to the north, the Atlantic Ocean to the east, Georgia and South Carolina to the south, and Tennessee to the west. Raleigh is the state's capital and Charlotte is its largest city. The Charlotte metropolitan area, with an estimated population of 2,569,213 in 2018, is the most populous metropolitan area in North Carolina, the 23rd-most populous in the United States, and the largest banking center in the nation after New York City. North Carolina's second largest metropolitan area is the Raleigh metropolitan area, with an estimated population of 1,337,331 in 2018, and is home to the largest research park in the United States, Research Triangle Park, in Chapel Hill, Durham, and Raleigh.

Florida is the southernmost contiguous state in the United States. The state is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Atlantic Ocean, and to the south by the Straits of Florida. Florida is the 22nd-most extensive, the 3rd-most populous, and the 8th-most densely populated of the U.S. states. Jacksonville is the most populous municipality in the state and the largest city by area in the contiguous United States. The Miami metropolitan area is Florida's most populous urban area. Tallahassee is the state's capital.

Ilex vomitoria, commonly known as yaupon or yaupon holly, is a species of holly that is native to southeastern North America. The word yaupon was derived from its Catawban name, yopún, which is a diminutive form of the word yop, meaning "tree". Another common name, cassina, was borrowed from Timucua. The Latin name comes from an incorrect belief by Europeans that the plant caused vomiting in certain ceremonies.


Related Research Articles

Parasitism is a symbiotic 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 characterised parasites as "predators that eat prey in units of less than one". Parasites include protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery animals such as hookworms, lice, mosquitoes, and vampire bats fungi such as honey fungus and the agents of ringworm and plants such as mistletoe, dodder, and the broomrapes. There are six major parasitic strategies of exploitation of animal hosts, namely parasitic castration, directly transmitted parasitism, trophically transmitted parasitism, vector-transmitted parasitism, parasitoidism, and micropredation.

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 pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in social insects such as ants, wasps, bees, and termites.

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

The Apocrita are a suborder of insects in the order Hymenoptera. It includes wasps, bees, and ants, and consists of many families. It contains the most advanced hymenopterans and is distinguished from Symphyta by the narrow "waist" (petiole) formed between the first two segments of the actual abdomen the first abdominal segment is fused to the thorax, and is called the propodeum. Therefore, it is general practice, when discussing the body of an apocritan in a technical sense, to refer to the mesosoma and metasoma rather than the "thorax" and "abdomen", respectively. The evolution of a constricted waist was an important adaption for the parasitoid lifestyle of the ancestral apocritan, allowing more maneuverability of the female's ovipositor. The ovipositor either extends freely or is retracted, and may be developed into a stinger for both defense and paralyzing prey. Larvae are legless and blind, and either feed inside a host or in a nest cell provisioned by their mothers.

Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causing the death of these hosts. Different species specialise in hosts from different insect orders, most often Lepidoptera, though some select beetles, flies, or bugs the spider wasps (Pompilidae) exclusively attack spiders.

The Perilampidae are a small family within the Chalcidoidea, composed mostly of hyperparasitoids. The family is closely related to the Eucharitidae, and the eucharitids appear to have evolved from within the Perilampidae, thus rendering the family paraphyletic. As presently defined, at least 15 genera and 270 species are described worldwide. They are often brilliantly metallic, with robust mesosomae and a small, triangular metasomae. They are generally very strongly sculptured. The prothorax is typically very broad and disc-like, and the labrum is multidigitate, a feature shared with the Eucharitidae.

The Leucospidae are a specialized group of wasps within the superfamily Chalcidoidea, that are ectoparasitoids of aculeate wasps or bees. They are typically mimics of bees or stinging wasps, often black with yellow, red, or white markings, sometimes metallic, with a robust mesosoma and very strong sculpturing. The hind femora are often greatly enlarged, with a row of teeth or serrations along the lower margin as in Chalcididae. The wing has a longitudinal fold. The female ovipositor is sometimes short, but if not, it is recurved and lies along the dorsal side of the metasoma, a unique feature. The males are also unusual, in the fusion of many of the metasomal segments to form a capsule-like "carapace".

Jalmenus evagoras, the imperial hairstreak, imperial blue, or common imperial blue, is a small, metallic blue butterfly of the family Lycaenidae. It is commonly found in eastern coastal regions of Australia. This species is notable for its unique mutualism with ants of the genus Iridomyrmex. The ants provide protection for juveniles and cues for adult mating behavior. They are compensated with food secreted from J. evagoras larvae. The ants greatly enhance the survival and reproductive success of the butterflies. J. evagoras lives and feeds on Acacia plants, so butterfly populations are localized to areas with preferred species of both host plants and ants.

Bombus bohemicus, also known as the gypsy's cuckoo bumblebee, is a species of socially parasitic cuckoo bumblebee found in most of Europe with the exception of the southern Iberian Peninsula and Iceland. B. bohemicus practices inquilinism, or brood parasitism, of other bumblebee species. B. bohemicus is a generalist parasite, successfully invading several species from genus Bombus. The invading queen mimics the host nest's chemical signals, allowing her to assume a reproductively dominant role as well as manipulation of host worker fertility and behavior.

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. The wasps do not constitute a clade, a complete natural group with a single ancestor, as their common ancestor is shared by bees and ants. Many wasps, those in the clade Aculeata, can sting their insect prey.

Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes metricus can be found attached to the sides of buildings, trees, and shrubbery.

Dolichovespula adulterina is a species of parasitic social wasp. D. adulterina lives in the Palearctic and Nearctic regions but parasitise different host species depending on which region it inhabits. D. adulterina feeds on a variety of foods including insects, spiders, arthropods, meat, molluscs, fruit, nectar and larval secretions. D. adulterina is synonymous with D. arctica from the Palearctic region.

Apocrypta is an Old World genus of parasitic fig wasps in the family Pteromalidae. They are parasitoids of gall-wasps in the Sycophagini tribe, and especially Ceratosolen species, pollinators of the Sycomorus, Sycocarpus and Neomorphe sections of Ficus. They seem to be fig species-specific.

Polistes biglumis is a species of social wasp within Polistes, the most common genus of paper wasp. It is distinguished mainly by its tendency to reside in montane climates in meadows or alpine areas. Selection pressure from the wasp's environment has led to several idiosyncrasies of its behavior and life cycle with respect to its relative species in the genus Polistes. It alone among paper wasps is often polyandrous. In addition, it has a truncated nesting season that gives rise to unique competitive dynamics among females of the species. P. biglumis wasps utilize an odor based recognition system that is the basis for all wasp to wasp interaction of the species. The wasp's life cycle is highly intertwined with that of Polistes atrimandibularis, an obligate social parasite wasp that frequently invades the combs of P. biglumis wasps.

Polistes sulcifer is a species of paper wasp in the genus Polistes that is found in Italy and Croatia. It is one of only three known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. sulcifer females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.

Latina is a genus of South American chalcid wasps in the family Eucharitidae. There are four known species of Latina with three known in Argentina and one from Venezuela.

Tamarixia radiata, the Asian citrus cyllid parasitoid, is an hymenopteran wasp from the family Eulophidae which was discovered in the 1920s in the area of northwestern India (Punjab), now Pakistan. It is a parasitoid of the Asian citrus psyllid, an economically important pest of citrus crops around the world and a vector for Citrus greening disease.

Melittobia australica is a species of chalcid wasp from the family Eulophidae which is a gregarious ecto-parasitoid of acuealate Hymenoptera.

Oraseminae is a subfamily of chalcid wasps in the family Eucharitidae. There are at least 10 genera in Oraseminae.

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.


Watch the video: Klecanka polna Polistes nimpha (November 2022).