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All hymenopterans (bees, ants, wasps, and sawflies) have haplodiploid sex determination. Generally, this involves haploid males developing from unfertilized eggs and diploid females developing from fertilized eggs, but diploid male production (DMP) frequently occurs. Some species have complementary sex determination (CSD), in which heterozygotes of a csd locus or loci are diploid females, hemizygotes are haploid males, and homozygotes are typically sterile diploid males. These diploid males underlie the diploid male vortex (DMV), the phenomenon of inbreeding reducing csd allelic diversity and causing increasingly higher production of sterile diploid males until the population dies out. The DMV has been cited as an extinction risk for many species across the order and a danger to both human-controlled and natural hymenopteran populations important in biological control, pollination, and endangered species conservation. However, it has been unclear how frequently it occurs. We review the known mechanisms for DMV avoidance. Many of these mechanisms are linked to lifestyle, and so we structure our investigation around eusocial lifestyle and non-eusocial (solitary or gregarious) species. We also review documented DMV incidence to make an inference about its prevalence. There are many means to avoid inbreeding or diploid male production, including eusocial-exclusive polygyny, polyandry, and diploid queen execution. For both eusocial and non-eusocial species these include biological mechanisms such as multi-locus CSD or non-CSD sex determination, diploid male inviability or brood removal, partial diploid male fecundity, and mating avoidance of kin or diploid males; or population genetic mechanisms such as gene flow and dispersal, balancing selection of csd alleles (with some being more prominent in eusocial vs. non-eusocial species). Documented cases of DMV are uncommon, and incidence is often tied to exacerbatory conditions such as habitat fragmentation and host declines. With this review we suggest that due to numerous avoidance mechanisms, DMV risk may not be as high as previously believed, and requires specific circumstances. In doing so we aim to better inform hymenopteran breeding and conservation efforts.

All hymenopterans (bees, ants, wasps, and sawflies) have haplodiploid sex determination. Generally, this involves haploid males developing from unfertilized eggs and diploid females developing from fertilized eggs, but diploid male production (DMP) frequently occurs. Some species have complementary sex determination (CSD), in which heterozygotes of a csd locus or loci are diploid females, hemizygotes are haploid males, and homozygotes are typically sterile diploid males. These diploid males underlie the diploid male vortex (DMV), the phenomenon of inbreeding reducing csd allelic diversity and causing increasingly higher production of sterile diploid males until the population dies out. The DMV has been cited as an extinction risk for many species across the order and a danger to both human-controlled and natural hymenopteran populations important in biological control, pollination, and endangered species conservation. However, it has been unclear how frequently it occurs. We review the known mechanisms for DMV avoidance. Many of these mechanisms are linked to lifestyle, and so we structure our investigation around eusocial lifestyle and non-eusocial (solitary or gregarious) species. We also review documented DMV incidence to make an inference about its prevalence. There are many means to avoid inbreeding or diploid male production, including eusocial-exclusive polygyny, polyandry, and diploid queen execution. For both eusocial and non-eusocial species these include biological mechanisms such as multi-locus CSD or non-CSD sex determination, diploid male inviability or brood removal, partial diploid male fecundity, and mating avoidance of kin or diploid males; or population genetic mechanisms such as gene flow and dispersal, balancing selection of csd alleles (with some being more prominent in eusocial vs. non-eusocial species). Documented cases of DMV are uncommon, and incidence is often tied to exacerbatory conditions such as habitat fragmentation and host declines. With this review we suggest that due to numerous avoidance mechanisms, DMV risk may not be as high as previously believed, and requires specific circumstances. In doing so we aim to better inform hymenopteran breeding and conservation efforts.

In insect sex determination a primary signal starts the genetic sex determination cascade that, in most insect orders, is subsequently transduced down the cascade by a transformer (tra) ortholog. Only a female-specifically spliced tra mRNA yields a functional TRA-protein that forms a complex with TRA2, encoded by a transformer-2 (tra2) ortholog, to act as a sex specific splicing regulator of the downstream transcription factors doublesex (dsx) and fruitless (fru). Here, we identify the tra2 ortholog of the haplodiploid parasitoid wasp N. vitripennis (Nv-tra2) and confirm its function in N. vitripennis sex determination. Knock down of Nv-tra2 by parental RNA interference (pRNAi) results in complete sex reversal of diploid offspring from female to male, indicating the requirement of Nv-tra2 for female sex determination. As Nv-tra2 pRNAi leads to frequent lethality in early developmental stages, maternal provision of Nv-tra2 transcripts is apparently also required for another, non-sex determining function during embryogenesis. In addition, lethality following Nv-tra2 pRNAi appears more pronounced in diploid than in haploid offspring. This diploid lethal effect was also observed following Nv-tra pRNAi, which served as a positive control in our experiments. As diploid embryos from fertilized eggs have a paternal chromosome set in addition to the maternal one, this suggests that either the presence of this paternal chromosome set or the dosage effect resulting from the diploid state is incompatible with the induced male development in N. vitripennis caused by either Nv-tra2 or Nv-tra pRNAi. The role of Nv-tra2 in activating the female sex determination pathway yields more insight into the sex determination mechanism of Nasonia. (C) 2017 The Authors. Published by Elsevier Ltd.