![141468-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-06/141468-Thumbnail%20Image.png?versionId=Uswnd7ikEp1KgDARcL3iblJ1UJk2.xqa&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240605/us-west-2/s3/aws4_request&X-Amz-Date=20240605T151840Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=982ede38fb04655146012bdf209b6d8f10d4d5ff5481ba9aed9b01ee7afedc7b&itok=x5O3xTiu)
In this synthesis, we hope to accomplish two things: 1) reflect on how the analysis of the new archaeological cases presented in this special feature adds to previous case studies by revisiting a set of propositions reported in a 2006 special feature, and 2) reflect on four main ideas that are more specific to the archaeological cases: i) societal choices are influenced by robustness–vulnerability trade-offs, ii) there is interplay between robustness–vulnerability trade-offs and robustness–performance trade-offs, iii) societies often get locked in to particular strategies, and iv) multiple positive feedbacks escalate the perceived cost of societal change. We then discuss whether these lock-in traps can be prevented or whether the risks associated with them can be mitigated. We conclude by highlighting how these long-term historical studies can help us to understand current society, societal practices, and the nexus between ecology and society.
![130905-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-05/130905-Thumbnail%20Image.png?versionId=Qd3k.RqPsAfqvacx8xMCUMn7QKAYDTnl&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240616/us-west-2/s3/aws4_request&X-Amz-Date=20240616T023325Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=6fcb10b15da91aef7dd0b9f336427fca473e2ce40cff9c7fcd7ed74290478fe5&itok=Am7jBq4Q)
![130412-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-04/130412-Thumbnail%20Image.png?versionId=6qY4LuypXX9EeorWO5uK_zKaCk3U9.pG&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240608/us-west-2/s3/aws4_request&X-Amz-Date=20240608T110551Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=7851a04c38e615d60199bcb23c864bd4d0767545bc430522b147dd7adb9d6880&itok=xDaihN5G)
![132764-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-05/132764-Thumbnail%20Image.png?versionId=jhVDQvexr9mOFGuWt4W5UYZDayMIr9oq&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240616/us-west-2/s3/aws4_request&X-Amz-Date=20240616T030350Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=b5c6374d497cf8543a00d830bc23fb5c1f5908a64e9412cbb1b904cf83fc204a&itok=Cv0KYOJB)
![132662-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-05/132662-Thumbnail%20Image.png?versionId=1TqK1h.kZCYZ9CSwiMxGKpSD1FZ83reO&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240615/us-west-2/s3/aws4_request&X-Amz-Date=20240615T081025Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=ec523be895383fc2648c3f34384ae3a31700e42e2aa992d01da71c238024232d&itok=C0IsbCbP)
![132078-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-08/132078-Thumbnail%20Image.png?versionId=XaueuUGqZfXBA1G9DcoBqpS5.Eyrq9B4&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240615/us-west-2/s3/aws4_request&X-Amz-Date=20240615T130354Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=f2e19d7b28bded460cbf1c2035a02283f1de09d1990b4beaaf0bab228c13a636&itok=vafu9N-y)
![130355-Thumbnail Image.png](https://d1rbsgppyrdqq4.cloudfront.net/s3fs-public/styles/width_400/public/2021-04/130355-Thumbnail%20Image.png?versionId=EBj.AazihhMn.qGGliOQ8N6UZqugxYwh&X-Amz-Content-Sha256=UNSIGNED-PAYLOAD&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Credential=AKIASBVQ3ZQ42ZLA5CUJ/20240605/us-west-2/s3/aws4_request&X-Amz-Date=20240605T191133Z&X-Amz-SignedHeaders=host&X-Amz-Expires=120&X-Amz-Signature=ec5d46192dc5d7c4ab640bf26c68d7e351a1ffd38f3dc5ec211c8e191251a438&itok=W3F4Jba7)
The reproductive ground plan hypothesis of social evolution suggests that reproductive controls of a solitary ancestor have been co-opted during social evolution, facilitating the division of labor among social insect workers. Despite substantial empirical support, the generality of this hypothesis is not universally accepted. Thus, we investigated the prediction of particular genes with pleiotropic effects on ovarian traits and social behavior in worker honey bees as a stringent test of the reproductive ground plan hypothesis. We complemented these tests with a comprehensive genome scan for additional quantitative trait loci (QTL) to gain a better understanding of the genetic architecture of the ovary size of honey bee workers, a morphological trait that is significant for understanding social insect caste evolution and general insect biology.
Results
Back-crossing hybrid European x Africanized honey bee queens to the Africanized parent colony generated two study populations with extraordinarily large worker ovaries. Despite the transgressive ovary phenotypes, several previously mapped QTL for social foraging behavior demonstrated ovary size effects, confirming the prediction of pleiotropic genetic effects on reproductive traits and social behavior. One major QTL for ovary size was detected in each backcross, along with several smaller effects and two QTL for ovary asymmetry. One of the main ovary size QTL coincided with a major QTL for ovary activation, explaining 3/4 of the phenotypic variance, although no simple positive correlation between ovary size and activation was observed.
Conclusions
Our results provide strong support for the reproductive ground plan hypothesis of evolution in study populations that are independent of the genetic stocks that originally led to the formulation of this hypothesis. As predicted, worker ovary size is genetically linked to multiple correlated traits of the complex division of labor in worker honey bees, known as the pollen hoarding syndrome. The genetic architecture of worker ovary size presumably consists of a combination of trait-specific loci and general regulators that affect the whole behavioral syndrome and may even play a role in caste determination. Several promising candidate genes in the QTL intervals await further study to clarify their potential role in social insect evolution and the regulation of insect fertility in general.