Only more recently has it become clear that many eusocial species also regularly reproduce. In the termite cavitermes tuberosus, parthenogenesis is an integral part of the lifecycle. Queens are adapted for reproduction while workers are adapted for foraging and colony maintenance. Thelytokous parthenogenesis in the fungusgardening ant. Ants evolved from wasplike ancestors in the midcretaceous period between 110 and million years ago and diversified after the rise of flowering plants. Induction of a thelytokous reproduction in the aphidius. Thelytokous parthenogenesis by queens in the dacetine ant pyramica membranifera hymenoptera. Taxonomic papers describing single species are acceptable if the species has economic importance or provides new data on the biology or evolution of the genus or higher. Additional information there are other files related to this item in huscap. Highly eusocial insect societies, such as all known ants, are typically characterized by a reproductive division of labor between queens, who are inseminated and reproduce, and virgin workers, who engage in foraging, nest maintenance and brood care.
Herbivory is common among the primitive hymenoptera suborder symphyta, in the gall wasps cynipidae, and in some of the ants and bees. Multiple mating and supercoloniality in cataglyphis desert ants. Loss of males from mixedsex societies in termites bmc. Honey bees and bumble bees associate with hostspecific lactobacilli, and some evidence suggests that these lactobacilli are. Eusocial hymenoptera ants, bees, wasps show remarkable variation in breeding. In eusocial hymenoptera ants, bees, wasps, at least three reproductive characteristics were shown to profoundly affect colony kin structure and may have played a key role in the evolution and maintenance of reproductive altruism. Apomictic parthenogenesis in a parasitoid wasp meteorus pulchricornis, uncommon in the haplodiploid order hymenoptera. Asexual reproduction in introduced and native populations of the ant cerapachys biroi.
We did not detect significant preference by arrhenotokous. It was established in 1992, and transferred to publishing with pensoft publishers in 2011, under an open access system. This is in accordance with what is known for other thelytokous eusocial hymenoptera. Figitidae from japan, with description of two new species author. Two distinct asexual lineages from multiqueen colonies likely originated from the same sexual singlequeen population.
Because honey bee queens mate with and store sperm from multiple drones, emergency queen. Thelytokous parthenogenesis in eusocial hymenoptera, annual. So far, thelytoky has been convincingly demonstrated in the cape honeybee, apis mellifera capensis 1921, and for seven. Thelytokous parthenogenesis has resulted in the evolution of. Mycocepurus smithii is unique among ants and among eusocial hymenoptera, in that males seem to be completely absent and only queens and not workers produce diploid offspring via. In this thesis i investigate the evolutionary and genetic consequences of thelytokous parthenogenesis, with particular focus on the cape honey bee apis mellifera capensis. Due to this phenomenon, a queenless and broodless colony of cape bees, in any other race destined to perish, is capable of selfpreservation. Eusociality evolved repeatedly in different orders of animals, particularly the hymenoptera the wasps, bees, and ants. The cytological mechanism underlying thelytoky in cape honey bees is automixis with central fusion 112, which leads to gradual loss of heterozygosity, especially at loci distant from centromeres 3, 84.
Thelytokous parthenogenesis by queens in the dacetine ant. Within the formicidae, thelytoky has been demonstrated only in four phylogenetically very distant species, including p. A decade ago, the mixed reproductive strategy asexual queen succession aqs was first described in termites. Independent colony founding was successful for a few queens. In eusocial hymenoptera ants, bees, wasps, at least three reproductive characteristics were shown to profoundly affect colony kin structure and may have played a key role in the evolution and maintenance of reproductive.
Evolutionary history of the hymenoptera sciencedirect. Hymenoptera sawflies, wasps, ants, and bees are one of four megadiverse insect orders, comprising more than 153,000 described and possibly up to one million undescribed extant species 1, 2. To establish the clonal raider ant cerapachys biroi as a model eusocial organism, we sequenced and assembled its 214 megabase mb draft genome using 33 gigabases gb of illumina reads 119. Morphological queens, both alate and dealate, were present in only 29 percent of the colonies. Specificity between lactobacilli and hymenopteran hosts is. To date, the complete loss of males from advanced social animal lineages has been reported only in ants and honey bees hymenoptera, whose workers are always female and whose males display no helping behaviors even. Female parthenogenesis, or thelytoky, is particularly common in solitary hymenoptera. Populations of thrips tabaci are known to have two sympatric but genetically isolated reproductive modes, arrhenotoky sexual reproduction and thelytoky asexual reproduction. The ability to produce diploid offspring from unfertilized eggs by thelytokous parthenogenesis is rare among the hymenoptera and provides a challenge to the theories mentioned above that try to explain the maintenance of eusociality stern and foster 1997. According to estimates derived from the proposed phylogeny of the hymenoptera, eusociality has arisen independently in wasps, bees, and ants fig. According to the journal citation reports, the journal has a 20 impact factor of 0. It is more common in invertebrates, like arthropods, but it can. Hymenopterans, the membranewinged insects, include bees, ants, and a large number of other insect taxa collectively referred to as wasps. Mar 12, 2016 thelytokous parthenogenesis as described above generates embryos that are pseudoclones of their worker parent.
However, we show here that these specimens represent males of the congeneric species m. In september 2006 in stellenbosch, south africa, we reared a. Worker reproduction of the invasive yellow crazy ant. Apr 17, 2008 in thelytokous parthenogenesis by cape workers the central rather than the terminal or random pronuclei fuse to produce the restored diploid nucleus, as if one of the central maternal pronuclei takes the place of a sperm pronucleus.
Journal of hymenoptera research publishes papers of high scientific quality reporting comprehensive research on all aspects of hymenoptera, including biology, behavior, ecology, systematics, taxonomy, genetics, and morphology. Bacteriomeassociated wolbachia of the parthenogenetic. Thelytokous parthenogenesis is rare among animals and reported in about 1,500 species, about 1 in of described animal species, according to a 1984 study. In 16 colonies collected in florida, a total of 66 individuals differed morphologically from queens and workers. In this article we will discuss about parthenogenesis in insects. Thelytokous parthenogenesis does not appear to be a very common phenomenon in the hymenoptera. Seven qccs laid by parasites were homozygous at all loci. Thripidae, is an example in which sexual and asexual reproduction occur sympatrically 2224.
Widespread occurrence of asexual reproduction in higher. Thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. Thelytokous parthenogenesis was examined in artificial groups of virgin laboratoryreared workers, where one worker typically monopolized reproduction despite the presence of several individuals with elongated ovaries. Annals of the entomological society of america 816. Many eusocial insects are in one order of insects the hymenoptera wasps see left, bees, and ants. Thelytokous parthenogenesis, the production of diploid female offspring from unfertilized eggs, has been observed in a small number of eusocial hymenopteran species. Hymenoptera also happen to haplodiploid, which means that males are made from unfertilized eggs and thus carry one half of the maternal genotype only, whereas females are diploid derived half from mom and half from dad. Eusocial hymenoptera ants, bees, wasps show remarkable variation in breeding systems. Taxonomy and evolution of putative thelytokous species of leptopilina hymenoptera. Thelytokous parthenogenesis with central fusion, as occurs in a.
Colonies typically consist of a single oncemated queen, and her sterile worker offspring. Taxonomic papers describing single species are acceptable if the species has economic importance or provides new data. Multiple mating and supercoloniality in cataglyphis desert ants laurianne leniaud1, abraham heftez2, laurent grumiau1 and. Although the genus mischocyttarus polistinae is a large group of neotropical eusocial wasps, the cuticular hydrocarbons were only investigated in a few species. Cryptic royal subfamilies in honey bee apis mellifera. Midcoxal articulations and the phylogeny of the order hymenoptera. Thelytokous parthenogenesis is the production of females from unfertilized eggs. Polyploidy versus endosymbionts in obligately thelytokous. Worker reproduction of the invasive yellow crazy ant anoplolepis gracilipes chingchen lee 1,2, hirotaka nakao 3, shuping tseng 4,5, hungwei hsu 4, gwoli lin 4, jiawei tay 6, johan billen 7. Here we report for the first time on the occurrence of physogastric workers and apparent worker. Thelytokous and arrhenotokous reproduction in unmated queens. The origins of eusociality in hymenoptera the insects. Thelytokous parthenogenesis in eusocial hymenoptera annual.
Here we report for the first time on the occurrence of physogastric workers and apparent worker reproduction. Because males are produced by parthenogenesis they come from unfertilized eggs and thus only have one set of chromosomes, and females are produced from fertilized eggs, sisters from a singlymated mother share 75% of their. This choice is crucial as the queen is the sole reproductive in the colony and her quality is essential to its success. Summary laying workers of apis mellifera capensis escholtzproducing impatemate offspring are normal diploids with 2n 32 chromosomes. Cheating honeybee workers produce royal offspring europe.
It is patchily distributed and restricted to taxa at the distant tips of phylogenies. Reproductive division of labor is one of the key features of social insects. In many species, however, workers retain functional ovaries and can lay unfertilized male eggs or trophic eggs. Formicidae article pdf available in the science of nature 978. In haplodiploid insects, thelytoky can be encoded in their genomes, or induced by maternally inherited bacteria such. Thelytokous parthenogenesis in eusocial hymenoptera thelytokous parthenogenesis in eusocial hymenoptera rabeling, christian. If this is the first time you use this feature, you will be asked to authorise cambridge core to connect with your account. Thelytokous parthenogenesis in eusocial hymenoptera rabeling and kronauer 2 of the host colony 2, 5, 52, 57. The majority of endosymbiont induced thelytoky is automixis with gamete duplication 1, 6, except for bacterially facilitated apomictic thelytoky that was recorded in rickettsia infected neochrysocharis formosa. Thelytokous parthenogenesis can be encoded in arthropod genomes or can be induced by inherited microbial endosymbionts.
Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide a comprehensive. Thelytokous parthenogenesis as described above generates embryos that are pseudoclones of their worker parent. Thelytoky, the parthenogenetic development of females, has independently evolved in several insect orders yet the study of its mechanisms has so far mostly focussed on haplodiploid hymenoptera, while alternative mechanisms of thelytoky such as polyploidy are far less understood. Figitidae from japan, with description of two new species created date. In platythyrea punctata smith, 1858, queens, gamergates and parthenogenetic workers cooccur in the same species. Jun 16, 2010 thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. Asexual reproduction in introduced and native populations of. In basal primitively eusocial wasps, the role of cuticular hydrocarbons in the colonial recognition has been extensively studied in genus polistes polistinae and stenogastrinae. That is, thelytokous offspring receive one copy of the two grandparental genomes and are largely identical to their thelytokous parent, unless recombination during meiosis has resulted in loss of heterozygosity. The genome of the clonal raider ant cerapachys biroi. As parasitoids, predators, and pollinators, hymenoptera play a fundamental role in virtually all terrestrial ecosystems and are of substantial economic importance 1, 3. Thelytoky is associated with a number of fascinating and novel reproductive systems. Thelytokous parthenogenesis in unmated queen honeybees. Lactobacillaceae are well known for their roles in food fermentation, as probiotics, and in human health, but they can also be dominant members of the microbiota of some species of hymenoptera ants, bees, and wasps.
The hymenoptera is the only order besides the isoptera termites to have evolved complex social systems with division of labor. The evolutionary genetics of thelytokous parthenogenesis. Pdf thelytokous parthenogenesis in eusocial hymenoptera. Thelytokous parthenogenesis in unmated queen honeybees apis. In most species workers have little reproductive options left. The cuticular hydrocarbons profiles in the colonial. Detection of gene flow from sexual to asexual lineages in. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide. Colony structure and reproduction in the thelytokous. Workers can reproduce by both arrhenotokous and thelytokous parthenogenesis. In aqs species, the workers, soldiers and dispersing reproductives are produced through sexual reproduction, while nondispersing neotenic queens arise through automictic thelytokous parthenogenesis, replace the founding queen and mate with the founding king. Especially in ants, thelytoky underlies a variety of idiosyncratic life histories with unique evolutionary and ecological consequences. During emergency queen rearing, worker honey bees apis mellifera select several otherwise workerdestined larvae to instead rear as candidates to replace their dead or failing queen. Sexual reproduction is the norm in almost all animal species, and in many advanced animal societies, both males and females participate in social activities.
Usually an unfertilised ovum develops into a new individual only after the union with the sperm or fertilisation but in certain cases the development of the egg takes place without the. They usually produce haploid males by arrhenotokous parthenogenesis, both in the. Polyploidy versus endosymbionts in obligately thelytokous thrips. Hamilton proposed that eusociality arose in social hymenoptera by kin selection because of their interesting genetic sex determination trait of haplodiploidy. Multiple mating and supercoloniality in cataglyphis desert.
Mycocepurus smithii is unique among ants and among eusocial hymenoptera, in that males seem to be completely absent and only queens and not workers produce diploid offspring via thelytoky. Thelytokous parthenogenesis in eusocial hymenoptera. Research open access worker reproduction of the invasive yellow crazy ant anoplolepis gracilipes chingchen lee1,2, hirotaka nakao3, shuping tseng4,5, hungwei hsu4, gwoli lin4, jiawei tay6, johan billen7, fuminori ito3, chowyang lee8, chungchi lin1 and chincheng scotty yang5 abstract background. Although some populations have a classical haplodiploid reproduction system, in other populations queens are produced by thelytokous parthenogenesis, males are produced by a male clonality system. Joint evolution of asexuality and queen number in an ant. The hymenoptera include famous examples of social insects, such as honeybees and true ants. Thelytoky from the greek thelys female and tokos birth is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous definition of thelytokous by merriamwebster. Jan 07, 20 thelytokous parthenogenesis in eusocial hymenoptera thelytokous parthenogenesis in eusocial hymenoptera rabeling, christian. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide a comprehensive overview. Ants are social insects of the family formicidae f. This true sociality in animals, in which sterile individuals work to further the reproductive success of others, is found in termites, ambrosia beetles, galldwelling aphids, thrips, marine spongedwelling shrimp synalpheus regalis, naked molerats heterocephalus glaber.
To understand the diversification and key evolutionary transitions of hymenoptera, most notably from phytophagy to parasitoidism and predation and vice versa and from solitary to eusocial life, we inferred the phylogeny and divergence times of all major lineages of hymenoptera by analyzing 3,256 proteincoding genes in 173 insect species. Pdf thelytokous parthenogenesis by queens in the dacetine. Queens that emerged in the laboratory were kept with or without workers under laboratory conditions. Cerapachys biroi has the lowest loh rate of any thelytokous eusocial insect studied, losing less heterozygosity in 21 years mll1 than other species lose in a single generation. Here, we document thelytokous reproduction by queens in the polygynous species pyramica membranifera. Comparisons of life histories between living species with different degrees of social behavior.
Asexual reproduction in introduced and native populations. A queen may be reared from parthenogenetically produced female larvae, which will take over the colony after mating. Taxonomy and evolution of putative thelytokous species of. Jun 21, 2019 a decade ago, the mixed reproductive strategy asexual queen succession aqs was first described in termites. Herein, we report behavioral, ecological and genetic studies to determine whether there is gene flow between arrhenotokous and thelytokous t. Title taxonomy and evolution of putative thelytokous species of leptopilina hymenoptera. Thelytokous parthenogenesis in the fungusgardening ant mycocepurus smithii hymenoptera. An important evolutionary characteristic of the formicine subfamily ponerinae is the occurrence of various alternative reproductive tactics within single species.
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