Copidosoma floridanum

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Copidosoma floridanum
202112 Copidosoma floridanum.svg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Encyrtidae
Genus: Copidosoma
Species:
C. floridanum
Binomial name
Copidosoma floridanum
Ashmead, 1900
Synonyms
List
    • Berecyntus floridanusAshmead, 1900
    • Copidosoma brethesi(Blanchard, 1936)
    • Copidosoma daccaensis(Mani, 1941)
    • Copidosoma floridanus(Ashmead, 1900)
    • Copidosoma intermedium(Mercet, 1921)
    • Copidosoma japonicumAshmead, 1904
    • Copidosoma javae(Girault, 1917)
    • Copidosoma maculatum(Ishii, 1928)
    • Copidosoma phytometrae(Risbec, 1951)
    • Copidosoma walshi(Mercet, 1922)
    • Holcencyrtus calypso Crawford, 1914
    • Litomastix argentinusBrèthes, 1913
    • Litomastix brethesiBlanchard, 1936
    • Litomastix calypso(Crawford, 1914)
    • Litomastix daccaensisMani, 1941
    • Litomastix floridana(Ashmead, 1900)
    • Litomastix floridanus(Ashmead, 1900)
    • Litomastix intermediaMercet, 1921
    • Litomastix intermediusMercet, 1921
    • Litomastix javae(Girault, 1917)
    • Litomastix maculataIshii, 1928
    • Litomastix phytometrae(Risbec, 1951)
    • Litomastix walshiMercet, 1922

Copidosoma floridanum is a species of wasp in the family Encyrtidae which is primarily a parasitoid of moths in the subfamily Plusiinae. It has the largest recorded brood of any parasitoidal insect, at 3,055 individuals. [1] The life cycle begins when a female oviposits into the eggs of a suitable host species, laying one or two eggs per host. Each egg divides repeatedly and develops into a brood of multiple individuals, a phenomenon called polyembryony. The larvae grow inside their host, breaking free at the end of the host's own larval stage. [1]

Contents

A cosmopolitan species, Copidosoma floridanum is distributed worldwide. Because of its significance to agriculture as pest control and its phylogenic relationship with other important species, the wasp's genome is being sequenced by the Human Genome Sequencing Center as part of the i5K project, [2] which aims to sequence the genomes of 5,000 arthropods. [3]

Behavior

Reproductive altruism

As a putatively eusocial species, C. floridanum embodies only two of the four behavioral characteristics that characterize genuine eusociality: larvae live in groups, and there is reproductive division of labor, or reproductive altruism. [4] The second characteristic, reproductive altruism, is, in these wasps, manifested as a sterile soldier caste that has the sole purpose of protecting their reproductive clonal siblings throughout their larval stage. Reproductive altruism behavior plays a major role in the survival and reproductive success of C. floridanum. This species displays haplodiploid sex determination, which increases relatedness among females from 0.5 to 0.75 because males develop from unfertilized eggs and are therefore haploid while females develop from normally fertilized eggs and are therefore diploid. So, as a result of eusocial progeny allocation and a distinctive type of clonal development in parasitized hosts, polyembryonic wasps including C. floridanum are able to thrive. [5] [6] [7] Additionally, these wasps modify their caste ratios in response to interspecific competition, creating a trade-off between reproduction and defense, as the wasps adapt to the levels of competition within the group. [5]

Aggression and spite

C. floridanum produce eggs that divide clonally to produce larger broods. The polyembryonic wasp caste system consists of two separate groups: some of the embryos in a clone mature into reproductive larvae that ultimately develop into adults, while the other group consists of sterile soldier larvae that protect siblings from competitors. [8] At this ecological level, the soldiers' reproductive altruism is tied to clone-level allocation to defense; thus, in order to maximize the reproductive success of the siblings, soldiers risk their own chances of reproductive success (with no conflict between the soldiers themselves). [8] In his study, Giron argues that soldier aggression in this wasp species is inversely related to competitors' genetic relatedness, without respect to levels of resource competition. [8] In a later study, Giron sought to differentiate between the aggression of female and male soldiers, finding that the latter group is non-aggressive toward all competitors. [8]

Polyembryonic wasps, including C. floridanum, exhibit spite through instances of precocious larval development. [9] Spite provides an explanation for how natural selection can favor harmful behaviors that are costly to both the actor and the recipient; spite is typically considered a form of altruism that benefits a secondary recipient. [10] Two criteria demonstrate that spite is truly occurring: (i) the behavior is truly costly to the actor and does not provide a long-term direct benefit; and (ii) harming behaviors are directed toward relatively unrelated individuals. [10]

In C. floridanum, the process takes place in the following manner: the host insect lays two eggs in the eggs of moths, usually one male and one female, which proceed to divide asexually to produce a brood of clonal brothers and clonal sisters. [6] The wasp larvae then mature within the moth caterpillar, utilizing the moth as food throughout growth. Competition for resources limits how many adult wasps can emerge from the host; this indicates that negative relatedness likely exists within the brood. A portion of the larvae do not emerge, who serve as adults who forgo future reproduction in order to kill relatively unrelated opposite-sex siblings maturing in the same host before dying themselves; this special group of adult killers developed precociously. [11] Asymmetrical dispersal (defined as the sex differences in the scale of competition) and asymmetrical relatedness (brothers tend to be more related to sisters than the reverse) most likely serve as the evolutionary resolution of this conflict, in favor of the sisters. [11] This process, most importantly, frees up resources for closer relatives.

Soldier's spite behavior

C. floridanum gain interspecific competitive advantage over other competitors, including Glyptapanteles pallipes and Microplitis demolitor , primarily due to the presence of their soldier caste, whose fitness is limited to the survival of their clonal siblings. Uka studied the interspecific competition between C. floridanum male broods and G. pallipes in order to elucidate the defensive strategies of the former group. The C. floridanum progeny survival rate was greater than that of G. pallipes, regardless of the interval of oviposition. [12] C. floridanum gains a competitive advantage through its ability to physiologically suppress or putatively attack its adversaries. First off, they secrete a physiologically suppressive factor from in their labial glands as embryos or even in the tissue of the host and steadily supplied to the hemolymph. This toxic factor causes damage to G. pallipes larvae and ultimately leads to death. More specifically, C. floridanum induces a delay in competitor maturation, along with generating paralysis and weakness (Uki). Physical attack, on the other hand, has not been proven to be effective as a means of killing competitors. [12]

Cellular compatibility of wasp within host

The morula-stage embryo of C. floridanum invades the embryo of the host, utilizing adherent junctions to host cells. This is an effective evolutionary strategy, as other approaches could leave obvious wounds on the host cells, alerting competitors to the presence of this wasp species. As a result, these embryos can invade a phylogenetically distant host embryo (the moth) by taking advantage of the compatibility of its cells with host tissues. [13]

Kin discrimination

During larval development, an extraembryonic membrane surrounds each organism during maturation with the caterpillar host. A series of experiments performed by Giron and Strand proved that this membrane serves as the cue for kin selection; they demonstrated that attack rates were less common with kinship when the membrane was present. When the membrane was removed, attack rates between kin increased. [7] This study additionally demonstrated how the membrane functions as a cue for kin discrimination by switching the membranes between larvae. Researchers found that the soldiers were fooled into not killing relatively unrelated larvae that were encased by a transplanted membrane.

Caste-based identity: genetic and development influences

Recently, studies have been conducted to identify differentially expressed genes in C. floridanum castes that code for identifiable ions and proteins that the sterile soldiers, for instance, share. Soldiers and reproductive larvae express enzymes with the differential usage of proteinase inhibitors and ribosomal proteins. [14] More specifically, odorant binding proteins (OBPs) are utilized for kin recognition, along with the likely usage of toxin-like and SP genes in the ability of soldiers to murder competitors or as immune defense against potential pathogens. [14] The host's molting cycle plays a significant role in determining the identity of precocious and reproductive larvae. More specifically, the C. floridanum young mature in synchrony with specific phases within the moth's molting cycle. In the early stages of embryonic development, changes within the host's developmental program intrinsically influence caste determination. [15]

Related Research Articles

<span class="mw-page-title-main">Hymenoptera</span> Order of insects comprising sawflies, wasps, bees, and ants

Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. This ovipositor is often modified into a stinger. The young develop through holometabolism — that is, they have a wormlike larval stage and an inactive pupal stage before they reach adulthood.

<span class="mw-page-title-main">Chalcid wasp</span> Superfamily of wasps

Chalcid wasps are insects within the superfamily Chalcidoidea, part of the order Hymenoptera. The superfamily contains some 22,500 known species, and an estimated total diversity of more than 500,000 species, meaning the vast majority have yet to be discovered and described. The name "chalcid" is often confused with the name "chalcidid", though the latter refers strictly to one constituent family, the Chalcididae, rather than the superfamily as a whole; accordingly, most recent publications (e.g.,) use the name "chalcidoid" when referring to members of the superfamily.

<span class="mw-page-title-main">Parasitoid</span> Organism that lives with its host and kills it

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.

<span class="mw-page-title-main">Behavioral ecology</span> Study of the evolutionary basis for animal behavior due to ecological pressures

Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior?

<span class="mw-page-title-main">Ovoviviparity</span> Gestation type

Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop inside eggs that remain in the mother's body until they are ready to hatch.

<i>Polydnaviriformidae</i> Family of viruses

Polydnaviriformidae ( PDV) is a family of insect viriforms; members are known as polydnaviruses. There are two genera in the family: Bracoform and Ichnoviriform. Polydnaviruses form a symbiotic relationship with parasitoid wasps. Ichnoviriforms (IV) occur in Ichneumonid wasps and Bracoviriforms (BV) in Braconid wasps. The larvae of wasps in both of those groups are themselves parasitic on Lepidoptera, and the polydnaviruses are important in circumventing the immune response of their parasitized hosts. Little or no sequence homology exists between BV and IV, suggesting that the two genera have been evolving independently for a long time.

In fair division problems, spite is a phenomenon that occurs when a player's value of an allocation decreases when one or more other players' valuation increases. Thus, other things being equal, a player exhibiting spite will prefer an allocation in which other players receive less than more.

<span class="mw-page-title-main">Parasitoid wasp</span> Group of wasps

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.

<span class="mw-page-title-main">Encyrtidae</span> Family of wasps

Encyrtidae is a large family of parasitic wasps, with some 3710 described species in about 455 genera. The larvae of the majority are primary parasitoids on Hemiptera, though other hosts are attacked, and details of the life history can be variable. They are found throughout the world in virtually all habitats, and are extremely important as biological control agents. They may also present as an ecological threat to the population of some species. For example, the endangered Papilio homerus butterfly is parasitized at a rate of 77%, making them the main contributor to egg mortality in this butterfly species.

Polyembryony is the phenomenon of two or more embryos developing from a single fertilized egg. Due to the embryos resulting from the same egg, the embryos are identical to one another, but are genetically diverse from the parents. The genetic difference between the offspring and the parents, but the similarity among siblings, are significant distinctions between polyembryony and the process of budding and typical sexual reproduction. Polyembryony can occur in humans, resulting in identical twins, though the process is random and at a low frequency. Polyembryony occurs regularly in many species of vertebrates, invertebrates, and plants.

<span class="mw-page-title-main">Wasp</span> Group of insects

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 bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.

<span class="mw-page-title-main">Eusociality</span> Highest level of animal sociality a species can attain

Eusociality is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.

<i>Ardices canescens</i> Species of moth

Ardices canescens, the dark-spotted tiger moth or light ermine moth, is a moth in the family Erebidae that is found across most of Australia. It originally was included in the genus Spilosoma, but later the generic status of Ardices was proven.

<span class="mw-page-title-main">Evolution of eusociality</span> Origins of cooperative brood care

Eusociality evolved repeatedly in different orders of animals, notably termites and the Hymenoptera. This 'true sociality' in animals, in which sterile individuals work to further the reproductive success of others, is found in termites, ambrosia beetles, gall-dwelling aphids, thrips, marine sponge-dwelling shrimp, naked mole-rats, and many genera in the insect order Hymenoptera. The fact that eusociality has evolved so often in the Hymenoptera, but remains rare throughout the rest of the animal kingdom, has made its evolution a topic of debate among evolutionary biologists. Eusocial organisms at first appear to behave in stark contrast with simple interpretations of Darwinian evolution: passing on one's genes to the next generation, or fitness, is a central idea in evolutionary biology.

<i>Polistes metricus</i> Species of wasp

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.

<i>Polistes exclamans</i> Species of wasp

Polistes exclamans, the Guinea paper wasp, is a social wasp and is part of the family Vespidae of the order Hymenoptera. It is found throughout the United States, Mexico, the Bahamas, Jamaica and parts of Canada. Due to solitary nest founding by queens, P. exclamans has extended its range in the past few decades and now covers the eastern half of the United States, as well as part of the north. This expansion is typically attributed to changing global climate and temperatures. P. exclamans has three specific castes, including males, workers, and queens, but the dominance hierarchy is further distinguished by age. The older the wasp is, the higher it is in ranking within the colony. In most P. exclamans nests, there is one queen who lays all the eggs in the colony. The physiological similarities between the worker and queen castes have led to experiments attempting to distinguish the characteristics of these two castes and how they are determined, though males have easily identifiable physiological characteristics. Since P. exclamans live in relatively small, open combed nests, they are often subject to predators and parasites, such as Chalcoela iphitalis, Elasmus polistis, and birds. P. exclamans have defense and recognition strategies that help protect against these predators and parasites.

<i>Copidosoma</i> Genus of wasps

Copidosoma is a genus of chalcid wasps, which are parasitoids of Lepidoptera. The wasps are polyembryonic, depositing one or two eggs into their host which then develop into multiple offspring and have a soldier caste and a reproductive caste.

<i>Kladothrips</i> Genus of thrips

Kladothrips is a genus of Australian gall thrips. It is notable for including some of the few organisms outside of Hymenoptera that exhibit eusociality.

<i>Copidosoma koehleri</i> Species of insect

Copidosoma koehleri belongs to the family Encyrtidae and genus Copidosoma within the order Hymenoptera. It is a parasitoid and its hosts are mainly potato tuber moth but it also infest Andean potato tuber moth, Guatemalan potato tuber moth and Tomato leafminer. Super-parasitism is associated with C. koehleri.

<span class="mw-page-title-main">Brachys tesselatus</span> Species of beetle

Brachys tesselatus is a species of a leaf-mining beetle, primarily distributed along the southeast coast of the United States. Its distribution corresponds with the turkey oak, its main host plant. Although B. tesselatus primarily feeds and mines on turkey oak leaves, its impact on the host species is limited, allowing for the survival of both species.

References

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