Although this study was unable to resolve all the ambiguities in this clade. Here, we document thelytokous reproduction by queens in the polygynous species pyramica membranifera. Morphological queens, both alate and dealate, were present in only 29 percent of the colonies. The differential diagnosis of two eurytoma species that could be reared from asphodeline. Natural selection drives the evolution of ant life cycles pnas. 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. In platythyrea punctata smith, 1858, queens, gamergates and parthenogenetic workers cooccur in the same species. Well documented cases of thelytokous parthenogenesis in social insects ants, bees and termites, with details on the type of. 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. Finally, we consider the honey bee in the broader context of haplodiploid hymenoptera. Mymaridae, larval parasitoids of ophelimus maskelli ashmead hymenoptera. Superfamily trigonaloidea family trigonalidae superfamily ichneumonoidea family ichneumonidae family braconidae.
Taxonomy and evolution of putative thelytokous species of. This demonstrates once again how eusocial species can evolve to combine the benefits of both sexual and asexual reproduction in novel and. Asexual reproduction via thelytokous parthenogenesis is widespread in. In thelytokous species diploid females develop parthenogenetically from unfertilised eggs and there are no males 1, 2. This helps explain the abundance of eusocial genera within the order hymenoptera, including three separate origins within halcitid bees alone. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and h.
Tachysphex antillarumis one of five species of this. Thelytokous and arrhenotokous reproduction in unmated queens. It is found in several groups of hymenoptera, including apidae, aphelinidae, cynipidae, formicidae, ichneumonidae, and tenthredinidae. Thelytokous parthenogenesis by queens in the dacetine ant pyramica membranifera hymenoptera. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide. A range of finding have emerged in the last decade, such as. Influence of host tree on abundance of itoplectis conquisitor say hymenoptera. Engel, and others this clade has been studied by examining the mitochondrial dna. Thelytokous parthenogenesis in unmated queen honeybees. Two new species of phytophagous hymenoptera belonging to the families oryssidae and tenthredinidae, with notes on other sawflies. Figitidae from japan, with description of two new species author.
Pdf thelytokous parthenogenesis by queens in the dacetine. Hymenoptera ants, bees and wasps, as well as the diplo. However, there is considerable differentiation between species in extent and type of eusocial behavior. Inheritance of thelytoky in the honey bee apis mellifera capensis. Journal of hymenoptera research by international society of hymenopterists. 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. Details two new species of phytophagous hymenoptera. The higher eusocial hymenoptera are those social bees, wasps, and ants in which queens and workers differ morphologically. Review of the world genera of aphelinidae hymenoptera. Today, over 125 bcas are commercially available in europe weintraub and cheek, 2005. Eulophidae, also appeared recently in the middle east and mediterranean countries and is also spreading quickly in this area. Thelytokous parthenogenesis in eusocial hymenoptera. Especially in ants, thelytoky underlies a variety of idiosyncratic life histories with unique evolutionary and ecological consequences.
The phenomenon of thelytoky, the asexual production of diploid female offspring, is being increasingly uncovered among the eusocial hymenoptera. An important evolutionary characteristic of the formicine subfamily ponerinae is the occurrence of various alternative reproductive tactics within single species. Thelytokous parthenogenesis in eusocial hymenoptera annual. The distribution of thelytoky, arrhenotoky and androgenesis among.
Thelytokous parthenogenesis in unmated queen honeybees apis. Eurytomidae is a host of puklina asphodelinae hymenoptera. Aug 11, 2014 the genetic origin of advanced social organization has long been one of the outstanding problems of evolutionary biology. Two new australian species of stethynium hymenoptera. We argue that this is because species in which both queens and workers are thelytokous that are unlikely to persist over evolutionary time. The genus indabracon van achterberg hymenoptera, braconidae, braconinae in china, with description of four new species yang li, pu tang, xuexin chen 10. A queen may be reared from parthenogenetically produced female larvae, which will take over the colony after mating. The number of biological control agents bcas released in greenhouses has increased greatly. Apidae is an obligate, workerless bumble bee social parasite which invades nests of bombus affinis. Single locus complementary sex determination in hymenoptera. Due to this phenomenon, a queenless and broodless colony of cape bees, in any other race destined to perish, is capable of selfpreservation. Thelytoky is associated with a number of fascinating and novel reproductive systems.
Based on storks 1988 sampling of canopy trees in borneo, tropical hymenoptera, excluding ants, rival beetles as the most speciose insect order. Mycocepurus smithii is unique among ants and among eusocial hymenoptera, in that males seem to be completely absent and only queens and not workers. Publication date 1992topics hymenoptera periodicals publisher washington, d. Induction of a thelytokous reproduction in the aphidius. Author information1museum of comparative zoology labs, harvard university, cambridge, massachusetts 028, usa. The two species have several particular biological and ecological characteristic traits in common, like multivoltinous development, thelytokous parthenogenesis and host species range. Social hymenopterans appeared during the cretaceous. First records of gonatocerus dolichocerus ashmead, palaeoneura sp. Natural selection drives the evolution of ant life cycles. Hymenoptera originated in the triassic, with the oldest fossils belonging to the family xyelidae.
Summary laying workers of apis mellifera capensis escholtzproducing impatemate offspring are normal diploids with 2n 32 chromosomes. 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. Cytogenetic basis of thelytoky in apis mellifera capensis springerlink. Observations on the nesting behavior oftachysphex antillarum hymenoptera. It is more common in invertebrates, like arthropods, but it can.
Among the hymenopterans, ants are far and away the most social almost all species, most having highly refined labor division, and have even been noted. It can be induced in hymenoptera by the bacteria wolbachia and cardinium. The genetic origin of advanced social organization has long been one of the outstanding problems of evolutionary biology. Figitidae from japan, with description of two new species created date. So far, thelytoky has been convincingly demonstrated in the cape honeybee, apis mellifera capensis 1921, and for seven. Hymenoptera ants, bees, wasps, sawflies, horntails. Proceedings of the linnean society of new south wales.
Thelytokous parthenogenesis in eusocial hymenoptera thelytokous parthenogenesis in eusocial hymenoptera rabeling, christian. The origins of eusociality in hymenoptera the insects. Thelytokous parthenogenesis, the production of diploid female offspring from unfertilized eggs, has been observed in a small number of eusocial hymenopteran species. Herbivory is common among the primitive hymenoptera suborder symphyta, in the gall wasps cynipidae, and in some of the ants and bees. According to estimates derived from the proposed phylogeny of the hymenoptera, eusociality has arisen independently in wasps, bees, and ants fig. Queenworker conflict and social parasitism in bumble bees.
Molecular phylogeny and identification of the egyptian wasps. 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. Request pdf the distribution of thelytoky, arrhenotoky and androgenesis among castes in the eusocial hymenoptera thelytokous parthenogenesis is the. The insect order hymenoptera includes the ants, bees, wasps, and sawflies.
Independent colony founding was successful for a few queens. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide a comprehensive overview. Hamilton proposed that eusociality arose in social hymenoptera by kin selection because of their interesting genetic sex determination trait of haplodiploidy. 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.
Thelytoky can occur by different mechanisms, each of which has a different impact on the level of homozygosity. Hymenoptera families biology 291 with meuti at kenyon. Thelytokous parthenogenesis in the fungusgardening ant. Ichneumonidae, a polyphagous parasite of the european pine shoot moth, rhyacionia buoliana schiff. Female parthenogenesis, or thelytoky, is particularly common in solitary hymenoptera. Kronauer 20 thelytokous parthenogenesis in eusocial hymenoptera. A checklist of the families and superfamiles of the hymenoptera, with some links. Primitively eusocial hymenoptera species are monogamous, as pointed out by hughes et al. Data are still lacking, but closely related solitary species are under the same predationselection pressure, and they, too, can be expected to be monogamous. Thelytokous parthenogenesis in eusocial hymenoptera, annual.
Molecular phylogeny and identification of the egyptian. Jun 16, 2010 thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. As parasitoids, predators, and pollinators, hymenoptera play a fundamental role in virtually all terrestrial ecosystems and are of substantial economic importance 1, 3. Taxonomy and evolution of putative thelytokous species of leptopilina 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. Worker reproduction in the higher eusocial hymenoptera. The predominance of female sex is noted in the population. Only more recently has it become clear that many eusocial species also. Here we present an analysis of the major steps in ant evolution, based for the first time, to our knowledge, on combined recent advances in paleontology, phylogeny, and the study of contemporary life histories. Taxonomic papers describing single species are acceptable if the species has economic importance or provides new data.
In september 2006 in stellenbosch, south africa, we reared a. Journal of hymenoptera research connecting repositories. Data are presented on the typematerial representing the species described by linnaeus in his genera sphex, chrysis, vespa, apis and mutilla, the names here considered total 158. Pdf thelytoky, the asexual production of females, is rare in honey bees. Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants.
The chances of parthenogenesis are rare since aim st all females are fertilized prior to their emergence from the host pupa. Hymenopteran families order hymenoptera suborder symphyta. Over 150,000 living species of hymenoptera have been described, 2 3 in addition to over 2,000 extinct ones. Thelytokous parthenogenesis in eusocial hymenoptera antwiki. The monogamy hypothesis, formulated by jacobus boomsma in 2007, is currently the leading hypothesis concerning the initial evolution of eusociality in the hymenoptera.
This study investigated the genetic diversity among the three wasps, vespa orientalis linnaeus, polistes bucharensis erichson, and polistes mongolicus du buysson. Of the balance of 147, three are emendations and five are unavailable homonyms or names proposed in the synonymy of other species. The hymenoptera is the only order besides the isoptera termites to have evolved complex social systems with division of labor. Queens that emerged in the laboratory were kept with or without workers under laboratory conditions. Such differences extend to the reproductive system. In most faunas, they constitute about 20% of insect species. The hymenoptera is one of the vital and biggest insect orders comprising the bees, wasps, sawflies, and 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 parthenogenesis by queens in the dacetine ant. Pdf thelytokous parthenogenesis in eusocial hymenoptera. 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. Many eusocial insects are in one order of insects the hymenoptera wasps see left, bees, and ants.
One name is represented by a neotype specimen in lund, and one by a lectotype figure. Wasps are important to natural and biological pest control because they are predators or parasitoids of pest arthropods. Comparisons of life histories between living species with different degrees of social behavior. The evolution of this group has been intensively studied by alex rasnitsyn, michael s. A single gene causes thelytokous parthenogenesis, the defining.
Aphelinidae in french polynesia, with notes on egg parasitism of the glassywinged sharpshooter, homalodisca vitripennis germar hemiptera. This demonstrates once again how eusocial species can evolve to combine the benefits of both sexual and asexual reproduction in novel and unexpected ways. Subfamilies and tribes of the family aphelinidae hymenoptera. Holotype specimens in london, uppsala or stockholm represent 60 names. Only more recently has it become clear that many eusocial species also regularly reproduce. Evolutionary history of the hymenoptera sciencedirect.
Formicidae article pdf available in the science of nature 978. Thelytokous parthenogenesis in the fungusgardening ant mycocepurus smithii hymenoptera. We provide evidence of the causal forces of natural selection. Sep 02, 2014 the genetic origin of advanced social organization has long been one of the outstanding problems of evolutionary biology. 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. Classification of the parasitic hymenoptera of the family aphelinidae chalcidoidea. Colony structure and reproduction in the thelytokous. Fisher department of zoology, university of toronto, toronto, ontario, m5s 1a1, canada abstract. Thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. In 16 colonies collected in florida, a total of 66 individuals differed morphologically from queens and workers.
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