Thelytoky is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Eusociality from greek eu good and social, the highest level of organization of sociality, is defined by the following characteristics. Thelytokous parthenogenesis, or the ability to produce female offspring without mating, is rare in animals suomalainen et al. Thelytokous parthenogenesis in eusocial hymenoptera thelytokous parthenogenesis in eusocial hymenoptera rabeling, christian. Over 150,000 living species of hymenoptera have been described, in addition to over 2,000 extinct ones. Genetic reincarnation of workers as queens in the eastern. Thelytokous parthenogenesis in the fungusgardening ant mycocepurus smithii hymenoptera. Apomictic parthenogenesis in a parasitoid wasp meteorus pulchricornis, uncommon in the haplodiploid order hymenoptera. If high relatedness is the binding force of their clonal. Pdf thelytokous parthenogenesis in eusocial hymenoptera. In aphids alone, whose colonies comprise genetically identical members, eusocial behaviors have evolved independently on many occasions, possibly as many as 17 times because of apomictic parthenogenesis. Ants are eusocial insects of the family formicidae and, along with the related wasps and bees, belong to the order hymenoptera. Evolution of ageing, costs of reproduction and the. Furthermore, in a number of species, both the workers and queens do reproduce.
Thelytokous parthenogenesis in eusocial hymenoptera antwiki. In the ants, posterior abdominal mobility is increased even more by second and sometimes third constrictions between the second and third, and third and fourth, abdominal segments, which give rise to the. In many eusocial hymenoptera, a proportion of males are produced by workers. Thus, offspring are homozygous for a single maternal allele at all loci. Ants evolved from vespoid wasp ancestors in the cretaceous period, and diversified after the rise of. Thelytokous parthenogenesis in eusocial hymenoptera. The hymenoptera is the only order besides the isoptera termites to have evolved complex social systems with division of labor.
Eusocial insects provide special opportunities to elucidate the evolution of ageing as queens have apparently evaded costs of reproduction and reversed the fecunditylongevity tradeoff generally observed in nonsocial organisms. It is more common in invertebrates, like arthropods, but it can. Colonies typically consist of a single oncemated queen, and her sterile worker offspring. Observations of parthenogenesis in monitor lizards pdf. Stochastic simulation verifies that the obtained expression gives a. Thelytokous parthenogenesis by queens in the dacetine ant. Effective population size in eusocial hymenoptera with.
According to the wellsupported evolutionary theory of ageing, random extrinsic mortality through accidents or predation inevitably causes older cohorts to make smaller reproductive contributions to future generations. In contrast, thelytokous parthenogenesis, the production of female offspring from unfertilized eggs, has been rarely confirmed in eusocial hymenoptera, although the number of recognized. Thelytokous parthenogenesis in the exotic dacetine ant strumigenys rogeri hymenoptera. Ants appear in the fossil record across the globe in considerable diversity during the latest early cretaceous and early late cretaceous, suggesting an earlier origin. Sex ratio biases in termites provide evidence for kin.
Within eusocial hymenoptera, arrhenotokous parthenogenesis is a widely distributed phenomenon, and the reproductive modes in ants are particularly diverse. Thelytokous parthenogenesis is the production of females from unfertilized eggs. Rickettsia symbionts cause parthenogenetic reproduction in. Parthenogenesis is a natural form of asexual reproduction in which growth and development of. Author information1museum of comparative zoology labs, harvard university, cambridge, massachusetts 028, usa. Characterization and expression analysis of adipokinetic. Pdf thelytokous parthenogenesis in the fungusgardening. Herbivory is common among the primitive hymenoptera suborder symphyta, in the gall wasps cynipidae, and in some of the ants and bees. Behavioural castes, dominance and division of labour in a primitively eusocial wasp behavioural castes, dominance and division of labour in a primitively eusocial wasp chandrashekara, k gadagkar, raghavendra 19910112 00. Request permission export citation add to favorites track citation. Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. In honeybee queens this stimulus is the physical squeezing of the egg as it is laid s asaki and o baru 2002, so division occurs whether the egg is fertilized or not. Comparisons of life histories between living species with different degrees of social behavior. Nevertheless, since the sterile soldiers represent only a minority of the colony, all other individuals being fertile, the social aphids are considered as insects at an early stage of social evolution stern, 1998 when compared to the highly eusocial taxa, namely termites and eusocial hymenoptera.
Download pdf eusocial insects of the order hymenoptera display reproductive division of labour, whereby offspring are born to reproductive individuals but cared for by others workers in a. Only recently has it become clear that several species of eusocial hymenopterans regularly reproduce by. Thelytoky, the asexual production of females, is rare in honey bees. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and. Thelytoky also acts as an exaptation for the emergence of reproductive parasites, the most. Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide a comprehensive overview.
Only more recently has it become clear that many eusocial species also regularly reproduce thelytokously, and here we provide a comprehensive. Complex colonies have evolved in ants, bees, and wasps hymenoptera and in termites isoptera. Thelytokous parthenogenesis in the exotic dacetine ant strumigenys. If this is the first time you use this feature, you will be asked to authorise cambridge core to connect with your account.
Here, we document thelytokous reproduction by queens in the polygynous species pyramica membranifera. In the hymenoptera, however, an alternative mechanism is required because unfertilized eggs can develop by arrhenotokous parthenogenesis. In animals, parthenogenesis means development of an embryo from an unfertilized egg cell. Parthenogenesis, a reproductive strategy that involves development of a. Members of the hymenoptera are familiar to a general audience and common names exist for a large variety of groups. We discuss theories applied to explain eusocial traits in the isoptera and the hymenoptera in order to learn more about how and why eusociality evolved from subsocial ancestors. However, it is ubiquitous in workers of the cape honey bee apis mellifera capensis. 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. The relevant questions related to sexual selection, sperm competition and cryptic female choice in polyandrous eusocial hymenoptera therefore often differ from those addressed in non eusocial mating systems, but their unusual character may allow interesting experimental tests of the generality of sexual selection theory that cannot be performed. Only more recently has it become clear that many eusocial species also. Eusocial evolution in termites and hymenoptera springerlink.
So far, thelytoky has been convincingly demonstrated in the cape honeybee, apis mellifera capensis 1921, and for seven. Female parthenogenesis, or thelytoky, is particularly common in solitary hymenoptera. The genus rickettsia contains a group of obligate intracellular symbionts of eukaryotic cells and belongs to the family rickettsiaceae in the order rickettsiales of the alphaproteobacteria 58, 90. To assess the effect of male production by workers on the effective population size n e, a general expression of n e in hymenoptera with workerproduced males is derived on the basis of the genetic drift in the frequency of a neutral allele. They include greenwood borings the ambrosia beetle austroplatypus 41, plant galls aphids and thrips 42, 43, and, in the one known example. But how reproduction affects longevity in eusocial insects has rarely been tested experimentally. Over the past several decades, thelytokous parthenogenesis, the production of diploid female offspring by unmated females, has been observed in several species of eusocial hymenopterans. The wasp waist, contrary to many peoples initial expectations, is actually not located between the thorax and abdomen, but is a constriction between the first and second abdominal segments. The swine parthenote placentae often appears hypovascular. Mycocepurus smithii is unique among ants and among eusocial hymenoptera, in that males seem to be completely absent and only queens. Thelytoky allows some workers to be reincarnated into the queen phenotype, and thereby selects for reproductive competition among workers.
Queens that emerged in the laboratory were kept with or without workers under laboratory conditions. Thelytokous parthenogenesis, the production of diploid female offspring from unfertilized eggs, has been observed in a small number of eusocial hymenopteran species. Evolution of reproductive traits in cataglyphis desert. The origins of eusociality in hymenoptera the insects. Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. Especially in ants, thelytoky underlies a variety of idiosyncratic life histories with unique evolutionary and ecological consequences. However, in the hymenopteran insects, all of which are haplodiploid, thelytoky is widespread, having evolved at least 255 times normark, 2003. However, diplodiploid eusocial organisms for example, termites 31, ambrosia beetles 32, shrimp 33, and naked mole rats 34 or clonal eusocial organisms for example, aphids 35 lack. In the insect order hymenoptera which includes bees, wasps, and. 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. In many hymenopteran insects such as honeybees, female eggs are produced. Cataglyphis desert ants display unique variation in their breeding systems, making this genus a particularly interesting model to study the evolution of derived reproductive traits in eusocial hymenoptera.
Clonal insects that exhibit eusocial behavior include aphids and polyembryonic wasps. Colonies may be headed by a single or several queens, and queens may be singly or multiply mated. Many species have medical importance as they are pathogens of humans and other vertebrates. Thelytokous parthenogenesis in which diploid females are produced from unfertilized eggs, was recently reported for some ant species. 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. 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. Thelytokous parthenogenesis in unmated queen honeybees. Matsuura, in encyclopedia of animal behavior, 2010. Thelytokous parthenogenesis in the fungusgardening ant. Download fulltext pdf download fulltext pdf download fulltext pdf. Kin selection predicts femalebiased sex ratios in haplodiploid eusocial insects because sisters are more related to each other than to their brothers. In contrast to eusocial hymenoptera, thelytoky in termites, such as reticulitermes speratus and r. The evolution of ageing, costs of reproduction and life history represent central, interlinked topics in evolutionary ecology.