Disparity by caste and tribe: Understanding women's empowerment and health outcomes in India.

AIM: Women's empowerment status varies greatly in India according to caste, class, ethnicity and region. This paper aims to investigate the caste/tribe disparity in women's empowerment by region, the main correlates of each domain of empowerment, and the association of women's empowerment with nutritional and health care access outcomes, specifically anaemia, menstrual hygiene, and institutional delivery. METHODOLOGY: Using National Family Health Survey-5 (2019-2021) data, we have created a modified survey-based women's empowerment index (SWPER) using principal component analysis with Oblique varimax rotation. The first four components are interpreted as an attitude to violence, freedom of movement, decision-making power and social independence. Several multivariate regression models were used to understand the factor associated with empowerment and the association of women's empowerment with different health outcomes. RESULTS: The results indicate that women from the forward castes are the most empowered in most domains except decision-making. However, after controlling other background variables, the forward castes women are found to be the most empowered in attitude to violence, whereas Scheduled Castes and Scheduled Tribes women were found to be the most empowered women in decision-making. With regards to social independence, deprived castes women are more empowered than the forward castes women. The likelihood of empowerment in social independence domain increases with increasing wealth. There are wide regional variations in empowerment level between different social groups. Caste/tribe identity plays a significant role in determining health outcomes in India. Among all empowerment domains, social independence emerges as the most significant associated factor with improved health across all caste/tribe groups. CONCLUSION: The path to women's empowerment in India must recognize the intersectionality of caste/tribe identities, and address regional disparities. Social independence emerges as a critical determinant across all caste/tribe groups for improving health. Measures should be taken to empower women through the underlying factors of social independence.

Major changes in domain arrangements are associated with the evolution of termites.

Domains as functional protein units and their rearrangements along the phylogeny can shed light on the functional changes of proteomes associated with the evolution of complex traits like eusociality. This complex trait is associated with sterile soldiers and workers, and long-lived, highly fecund reproductives. Unlike in Hymenoptera (ants, bees, and wasps), the evolution of eusociality within Blattodea, where termites evolved from within cockroaches, was accompanied by a reduction in proteome size, raising the question of whether functional novelty was achieved with existing rather than novel proteins. To address this, we investigated the role of domain rearrangements during the evolution of termite eusociality. Analysing domain rearrangements in the proteomes of three solitary cockroaches and five eusocial termites, we inferred more than 5,000 rearrangements over the phylogeny of Blattodea. The 90 novel domain arrangements that emerged at the origin of termites were enriched for several functions related to longevity, such as protein homeostasis, DNA repair, mitochondrial activity, and nutrient sensing. Many domain rearrangements were related to changes in developmental pathways, important for the emergence of novel castes. Along with the elaboration of social complexity, including permanently sterile workers and larger, foraging colonies, we found 110 further domain arrangements with functions related to protein glycosylation and ion transport. We found an enrichment of caste-biased expression and splicing within rearranged genes, highlighting their importance for the evolution of castes. Furthermore, we found increased levels of DNA methylation among rearranged compared to non-rearranged genes suggesting fundamental differences in their regulation. Our findings indicate the importance of domain rearrangements in the generation of functional novelty necessary for termite eusociality to evolve.

Peeking into the Stingers: A Comprehensive SWATH-MS Study of the European Hornet Vespa crabro (Linnaeus, 1758) (Hymenoptera: Vespidae) Venom Sac Extracts.

This study aimed to investigate the venom sac extracts (VSEs) of the European hornet (EH) Vespa crabro (Linnaeus, 1758) (Hymenoptera: Vespidae), focusing on the differences between stinging females, gynes (G), and workers (W), at the protein level. Using a quantitative "Sequential Window Acquisition of all Theoretical Fragment Ion Mass Spectra" (SWATH-MS) analysis, we identified and quantified a total of 240 proteins. Notably, within the group, 45.8% (n = 110) showed significant differential expression between VSE-G and VSE-W. In this set, 57.3% (n = 63) were upregulated and 42.7% (n = 47) downregulated in the G. Additionally, the two-hundred quantified proteins from the class Insecta belong to sixteen different species, six of them to the Hymenoptera/Apidae lineage, comprising seven proteins with known potential allergenicity. Thus, phospholipase A1 (Vesp v 1), phospholipase A1 verutoxin 2b (VT-2b), hyaluronidase A (Vesp v 2A), hyaluronidase B (Vesp v 2B), and venom allergen 5 (Vesp v 5) were significantly downregulated in the G, and vitellogenin (Vesp v 6) was upregulated. Overall, 46% of the VSE proteins showed differential expression, with a majority being upregulated in G. Data are available via ProteomeXchange with identifier PXD047955. These findings shed light on the proteomic differences in VSE between EH castes, potentially contributing to our understanding of their behavior and offering insights for allergy research.

Novel insights into paternity skew in a polyandrous social wasp.

Females of many species are polyandrous. However, polyandry can give rise to conflict among individuals within families. We examined the level of polyandry and paternity skew in the common eastern yellowjacket wasp, Vespula maculifrons, in order to gain a greater understanding of conflict in social insects. We collected 10 colonies of V. maculifrons and genotyped workers and prereproductive queens at highly variable microsatellite markers to assign each to a patriline. Genotypic data revealed evidence of significant paternity skew among patrilines. In addition, we found that patrilines contributed differentially to caste production (worker vs. queen), suggesting an important role for reproductive conflict not previously discovered. We also investigated if patterns of paternity skew and mate number varied over time. However, we found no evidence of changes in levels of polyandry when compared to historical data dating back almost 40 years. Finally, we measured a suite of morphological traits in individuals from the most common and least common patrilines in each colony to test if males that showed highly skewed reproductive success also produced offspring that differed in phenotype. Our data revealed weak correlation between paternity skew and morphological phenotype of offspring sired by different males, suggesting no evidence of evolutionary tradeoffs at the level investigated. Overall, this study is the first to report significant paternity and caste-associated skew in V. maculifrons, and to investigate the phenotypic consequences of skew in a social wasp. Our results suggest that polyandry can have important consequences on the genetic and social structure of insect societies.

Handling heatwaves: balancing thermoregulation, foraging and bumblebee colony success.

Climate changes pose risks for bumblebee populations, which have declined relative to the growing frequency and severity of warmer temperature extremes. Bumblebees might mitigate the effects of such extreme weather through colonial behaviours. In particular, fanning behaviour to dissipate heat is an important mechanism that could reduce exposure of thermally sensitive offspring to detrimental nest temperatures (Tn). The allocation of workers towards fanning over prolonged periods could impact foraging activity that is essential for colony-sustaining resource gathering. Colony maintenance and growth could suffer as a result of nutritional and high ambient temperature (Ta) thermal stress. It remains uncertain whether a compromise occurs between thermoregulation and foraging under chronic, sublethal heat events and how colony success is impacted as a result. This study held colonies of Bombus impatiens at constant high Ta (25°C, 30°C or 35°C) for 2 weeks while quantifying the percentage of foragers, fanning incidence, nest temperature (Tn) and other metrics of colony success such as the percentage of adult emergence and offspring production. We found that foraging and adult emergence were not significantly affected by Ta, but that thermoregulation was unsuccessful at maintaining Tn despite increased fanning at 35°C. Furthermore, 35°C resulted in workers abandoning the colony and fewer offspring being produced. Our findings imply that heatwave events that exceed 30°C can negatively impact colony success through failed thermoregulation and reduced workforce production.

Age- and caste-independent piRNAs in the germline and miRNA profiles linked to caste and fecundity in the ant Temnothorax rugatulus.

Social insects are models for studies of phenotypic plasticity. Ant queens and workers vary in fecundity and lifespan, which are enhanced and extended in queens. Yet, the regulatory mechanisms underlying this variation are not well understood. Ant queens live and reproduce for years, so that they need to protect their germline from transposable element (TE) activity, which may be redundant in short-lived, often sterile workers. We analysed the expression of two protective classes of small RNAs, microRNAs (miRNAs) and Piwi-interacting RNAs (piRNAs), in various tissues, castes and age classes of the ant Temnothorax rugatulus. In queens, piRNAs were highly abundant in ovaries with TEs being their clear targets, with reduced but still detectable piRNA-specific ping-pong signatures in thorax and brains. piRNA pathway activity varied little with age in queens. Moreover, the reduced ovaries of workers also exhibited similar piRNA activity and this not only in young, fertile workers, but also in older foragers with regressed ovaries. Therefore, these ants protect their germline through piRNA activity, regardless of ovarian development, age or caste, even in sterile workers often considered the soma of the superorganism. Our tissue-specific miRNA analysis detected the expression of 304 miRNAs, of which 105 were expressed in all tissues, 10 enriched in the brain, three in the thorax, whereas 83 were ovarian-specific. We identified ovarian miRNAs whose expression was related to caste, fecundity and age, and which likely regulate group-specific gene expression. sRNA shifts in young- to middle-aged queens were minor, suggesting delayed senescence in this reproductive caste.

High temperature sensitivity of bumblebee castes and the colony-level costs of thermoregulation in Bombus impatiens.

Physiological thermal limits often reflect species distribution, but the role that ambient temperature (Ta) plays in limiting species within their thermal environment remains unclear. Climate change-linked declines in bumblebees, an important pollinator group, leave questions regarding which aspect of their physiology is hindered under high Ta. As a eusocial species, bumblebees utilize their ability to thermoregulate as a superorganism to maintain nest temperature (Tn) within a narrow thermal window to buffer developing larvae from developmental defects. Thermoregulatory behaviours, such as thermogenesis to warm up and fanning to cool down the nest, are energetically expensive and it is uncertain how successful large colonies are at maintaining Tn within its optimal range. Using a common bumblebee species, Bombus impatiens, our study first established the critical thermal limits (CTmax) of workers, queens, drones and larvae to determine which caste is most thermally sensitive to heat. We found that larvae had significantly lower heat tolerance than adults, highlighting the importance of colonial thermoregulation. We then measured the energy expenditure of large colonies under acute thermal stress (5-40 °C) using flow-through respirometry while simultaneously quantifying Tn. Colonies that experienced Ta at or below optimal Tn (≤30 °C) were successful at thermoregulation. At 35 °C and above, however, Tn increased despite high energetic costs to the colony. Together our results demonstrate that high Ta poses a risk to colonies that fail to buffer thermally sensitive larvae from changes in Tn.

Neuromodulators and neuroepigenetics of social behavior in ants.

Eusocial insects exemplify a remarkable system of division of labor within the same colony. This behavioral range, which is sometimes accompanied by morphological or physiological differences, provides an opportunity to study the relationship between complex behaviors and their underlying molecular mechanisms. This is especially true in ants because certain genera have an elaborate caste system and can dramatically change their stereotypical behavior over their lifetime. Recent studies experimentally alter ant behavior over short times, thus opening the study of underlying plasticity pathways. The molecular underpinnings of these behaviors are neuromodulators as well as the regulation of chromatin. Here, we concisely review the current understanding of the relationship between neuromodulators, epigenetics, and social behavior in ants. We discuss future directions in light of experimental limitations of the ant system.

Gene expression profiles of chemosensory genes of termite soldier and worker antennae.

Termites have an elaborate social system that involves cooperation and division of labour among colony members. Although this social system is regulated by chemical signals produced in the colony, it remains unclear how these signals are perceived by other members. Signal transduction is well known to be triggered by the reception of odorant molecules by some binding proteins in the antennae, after which, a signal is transmitted to chemosensory receptors. However, there is insufficient information on the role of chemosensory genes involved in signal transduction in termites. Here, we identified the genes involved in chemosensory reception in the termite Reticulitermes speratus and performed a genome-wide comparative transcriptome analysis of worker and soldier antennae. First, we identified 31 odorant-binding proteins (OBPs), and three chemosensory protein A (CheA) from the genome data. Thereafter, we performed RNA sequencing to compare the expression levels of OBPs, CheAs, and previously identified chemosensory receptor genes between worker and soldier antennae. There were no receptor genes with significant differences in expression between castes. However, the expression levels of three non-receptor odorant-detection/binding proteins (OBP, CheA, and Sensory neuron membrane protein) were significantly different between castes. Real-time qPCR (RT-qPCR) analysis using antennae and other head parts confirmed that these genes were highly expressed in soldier antennae. Finally, independent RT-qPCR analysis showed that the expression patterns of these genes were altered in soldiers from different social contexts. Present results suggest that gene expression levels of some non-receptors are affected by both castes and behavioural interactions among colony members in termites.

Comparison of Morphological Characteristics of Antennae and Antennal Sensilla among Four Species of Bumblebees (Hymenoptera: Apidae).

Bumblebees, as pollinators, play an important role in maintaining natural and agricultural ecosystems. Antennae with sensilla of bumblebees as social insects have essential effects in foraging, nest searching, courting, and mating, and are different in species and sexes. Previous studies on the morphology of antennae and sensilla in bumblebees have been limited to a few species and a single caste. To better understand how bumblebees detect and receive the chemical signal from nectariferous plants and foraging behavior, the morphology of antennae with sensilla, including the antennal length, and type, distribution, and number of antennal sensilla in four species, Bombus atripes, Bombus breviceps, Bombus flavescens, and Bombus terrestris was compared by scanning electron microscopy (SEM) herein. The total antennal length of queens are the longest and workers are the shortest in three castes, and in four species the longest of the total antennal length among three castes all are in B. flavescens, which is significantly longer than other species (p < 0.05) and the length of the scape in queens and workers are both longer than males, significantly different in queens (p < 0.05), and not significantly different in workers (p > 0.05), and the length of flagellums in females are not always shorter than males, of which the length of flagellms in queens of B. flavescens are significantly longer than males (p < 0.05), and the length of pedicel and all flagellomeres varies among species and castes. A total of 13 major types of sensilla in total were observed, including trichodea sensilla (TS A-E), placodea sensilla (PS A-B), basiconica sensilla (BaS), coeloconica sensilla (COS A-B), chaetic sensilla (CS A-B), and Böhm sensilla (BS), of which chaetic sensilla B (CS B), only observed in females of B. atripes, was firstly reported in Apidae. Moreover, the number of all sensilla was the most in males, the least was in workers, the number of sensilla varies within castes and species. Furthermore, the morphological characteristics of antennae and the potential functions of sensilla are discussed.

Co-expression Gene Networks and Machine-learning Algorithms Unveil a Core Genetic Toolkit for Reproductive Division of Labour in Rudimentary Insect Societies.

The evolution of eusociality requires that individuals forgo some or all their own reproduction to assist the reproduction of others in their group, such as a primary egg-laying queen. A major open question is how genes and genetic pathways sculpt the evolution of eusociality, especially in rudimentary forms of sociality-those with smaller cooperative nests when compared with species such as honeybees that possess large societies. We lack comprehensive comparative studies examining shared patterns and processes across multiple social lineages. Here we examine the mechanisms of molecular convergence across two lineages of bees and wasps exhibiting such rudimentary societies. These societies consist of few individuals and their life histories range from facultative to obligately social. Using six species across four independent origins of sociality, we conduct a comparative meta-analysis of publicly available transcriptomes. Standard methods detected little similarity in patterns of differential gene expression in brain transcriptomes among reproductive and non-reproductive individuals across species. By contrast, both supervised machine learning and consensus co-expression network approaches uncovered sets of genes with conserved expression patterns among reproductive and non-reproductive phenotypes across species. These sets overlap substantially, and may comprise a shared genetic "toolkit" for sociality across the distantly related taxa of bees and wasps and independently evolved lineages of sociality. We also found many lineage-specific genes and co-expression modules associated with social phenotypes and possible signatures of shared life-history traits. These results reveal how taxon-specific molecular mechanisms complement a core toolkit of molecular processes in sculpting traits related to the evolution of eusociality.

Association between parasite microbiomes and caste development and colony structure in a social trematode.

Division of labour through the formation of morphologically and functionally distinct castes is a recurring theme in the evolution of animal sociality. The mechanisms driving the differentiation of individuals into distinct castes remain poorly understood, especially for animals forming clonal colonies. We test the association between microbiomes and caste formation within the social trematode Philophthalmus attenuatus, using a metabarcoding approach targeting the bacterial 16S SSU rRNA gene. Clonal colonies of this trematode within snail hosts comprise large reproductive individuals which produce dispersal stages, and small, non-reproducing soldiers which defend the colony against invaders. In colonies extracted directly from field-collected snails, reproductives harboured more diverse bacterial communities than soldiers, and reproductives and soldiers harboured distinct bacterial communities, at all taxonomic levels considered. No single bacterial taxon showed high enough prevalence in either soldiers or reproductives to be singled out as a key driver, indicating that the whole microbial community contributes to these differences. Other colonies were experimentally exposed to antibiotics to alter their bacterial communities, and sampled shortly after treatment and weeks later after allowing for turnover of colony members. At those time points, bacterial communities of the two castes still differed across all antibiotic treatments; however, the caste ratio within colonies changed: after antibiotic disruption and turnover of individuals, new individuals were more likely to become reproductives than in undisturbed control colonies. Our results reveal that each caste has a distinct microbiome; whether the social context affects the microbiota, or whether microbes contribute to modulating the phenotype of individuals, remains to be determined.

Cuticular hydrocarbons as caste-linked cues in Neotropical swarm-founding wasps.

Wasps (Vespidae) are important organisms to understand the evolution of social behaviour. Wasps show different levels of sociality, which includes solitary to highly eusocial organisms. In social insect species, queens and workers differ in physiology and morphology. The Neotropical swarm-founding wasps (Epiponini) show a variety of caste syndromes. In this clade, the caste-flexibility is a unique characteristic, in which workers can become queens and swarm to start a new nest. The investigation of the caste system comparing several Epiponini species show a clear-cut morphological distinction between queens and workers, with a morphological continuum between queens and workers. However, whether cuticular hydrocarbons (CHCs) are used as cues for caste recognition in swarm-founding wasps is still unknown. We studied whether CHCs may display caste-linked differences in eleven species of Epiponini wasps and if CHCs differences would follow morphological patterns. Our results suggest that queens and workers of Epiponini wasps are chemically different from each other at two levels, qualitatively and quantitatively, or merely quantitatively. This variation seems to exist regardless of their morphological traits and may be useful to help us understanding how chemical communication evolved differently in these species.

Adverse childhood experiences and maternal self-efficacy: Examining the mediating role of intimate partner violence and the moderating role of caste membership in rural India.

Maternal self-efficacy (MSE) is associated with healthy functioning in mothers and children globally. Maternal exposure to adverse childhood experiences (ACEs) and intimate partner violence (IPV) is known to negatively impact MSE in high-income countries; however, the association has not been examined in low-and-middle-income countries, such as India, which face socioeconomic risks including poverty, illiteracy, and discrimination based on caste membership. The present study examines the mediating role of IPV in the association between ACEs (specifically-emotional, physical, and sexual abuse, neglect, household dysfunction, and discrimination) and MSE and tests caste membership as a moderator. A community-based, cross-sectional survey was performed with 316 mothers with at least one child between 0 and 24 months in a rural area in the North Indian state of Uttar Pradesh. A structural equation framework was used to test the moderated-mediation model. Results from the moderated-mediation model indicate that greater ACEs exposure was associated with lower MSE and this association was mediated by IPV exposure for low-caste but not high-caste mothers, even after controlling for wealth and literacy. These findings add to existing evidence on ACEs exposure as a significant burden for rural Indian mothers, negatively impacting parenting outcomes such as MSE. The critical role of caste membership is also highlighted, providing implications for future research.

La autoeficacia materna (MSE) se asocia con el saludable funcionamiento en las madres y niños globalmente. Se conoce que el hecho de que la madre haya estado expuesta a experiencias adversas en la niñez (ACE) y a la violencia de la pareja íntima (IPV) tiene un negativo impacto en MSE en países de altas entradas económicas; sin embargo, esta asociación no se ha examinado en países donde las entradas económicas son bajas o medias, como India, que enfrenta riesgos socioeconómicos entre los que se incluyen la pobreza, el analfabetismo, así como la discriminación basada en la pertenencia a una casta. El presente estudio examina el papel mediador de IPV en la asociación entre ACE (específicamente - el abuso emocional, físico y sexual, negligencia, disfuncionalidad en el hogar y discriminación) y MSE, y pone a prueba la pertenencia a una casta como aspecto moderador. Se llevó a cabo una encuesta de base comunitaria e inter-seccional con 316 madres con por lo menos un niño entre 0 y 24 meses de edad en un área rural en el estado de Uttar Pradesh en el norte de India. Se usó un marco de trabajo de ecuación estructural para examinar el modelo de moderación y mediación. Los resultados del modelo de moderación y mediación indican que una mayor exposición a ACE estaba asociada con una más baja MSE y que la exposición a IPV mediaba esa asociación para madres de castas bajas, pero no para madres de castas altas, aun después del factor control de recursos económicos y alfabetismo. Estos resultados contribuyen a la existente evidencia de que el haber estado expuesta a ACE es una carga significativa para las madres en la India rural, la cual tiene un impacto negativo en los resultados de crianza tales como MSE. También se subraya el papel esencial de la pertenencia a una casta, lo cual aporta implicaciones para la investigación futura.

L'auto-efficacité maternelle (MSE en anglais) est globalement liée à un fonctionnement sain chez les mères et les enfants. L'exposition maternelle à des expériences de l'enfance adverses (ACE en anglais) et à la violence intime ou conjugale (IPV) est connue comme impactant de manière négative l'auto-efficacité maternelle dans les pays à revenus élevés. Cependant ce lien n'a pas toujours été examiné dans les pays à faibles ou moyens revenus, tels que l'Inde qui fait face à des risques socioéconomiques qui comportent la pauvreté, l'illettrisme, la discrimination en fonction de l'appartenance à une caste. Cette étude examine le rôle médiateur de la violence conjugale (ou violence entre partenaires intimes) dans le lien entre les ACE (plus spécifiquement l'abus émotionnel, physiques, sexuel, la négligence, la dysfonction au sein du foyer et la discrimination) et l'auto-efficacité maternelle et les tests d'appartenance à une caste en tant que modérateurs. Un questionnaire communautaire, en coupe transversale, a été présenté à 116 mères ayant au moins un enfant entre l'âge de 0-24 mois dans une région rurale de l'état du nord de l'Inde, Uttar Pradesh. Un cadre d'équation structurelle a été utilisé pour tester le modèle de modération-médiation. Les résultats de ce modèle de modération-médiation indiquent que plus l'exposition aux ACE est grande, plus l'auto-efficacité maternelle est basse et cette association est affectée par l'exposition à la violence conjugale pour les castes moins élevées mais pas pour les mères des castes plus élevées, même en effectuant un contrôle pour la richesse et l'alphabétisation. Ces résultats s'ajoutent aux preuves existantes sur l'exposition aux ACE en tant que poids important pour les mères indienne de milieux ruraux, ce qui impacte de manière négative les résultats de parentage telles que l'auto-efficacité maternelle. Le rôle critique de l'appartenance à une caste est également mis en lumière, offrant des implications pour les recherches à venir.

Warm and arid regions of the world are hotspots of superorganism complexity.

Biologists have long been fascinated by the processes that give rise to phenotypic complexity of organisms, yet whether there exist geographical hotspots of phenotypic complexity remains poorly explored. Phenotypic complexity can be readily observed in ant colonies, which are superorganisms with morphologically differentiated queen and worker castes analogous to the germline and soma of multicellular organisms. Several ant species have evolved 'worker polymorphism', where workers in a single colony show quantifiable differences in size and head-to-body scaling. Here, we use 256 754 occurrence points from 8990 ant species to investigate the geography of worker polymorphism. We show that arid regions of the world are the hotspots of superorganism complexity. Tropical savannahs and deserts, which are typically species-poor relative to tropical or even temperate forests, harbour the highest densities of polymorphic ants. We discuss the possible adaptive advantages that worker polymorphism provides in arid environments. Our work may provide a window into the environmental conditions that promote the emergence of highly complex phenotypes.

Ecological Stoichiometry of Bumblebee Castes, Sexes, and Age Groups.

Ecological stoichiometry is important for revealing how the composition of chemical elements of organisms is influenced by their physiological functions and ecology. In this study, we investigated the elemental body composition of queens, workers, and males of the bumblebee Bombus terrestris, an important pollinator throughout Eurasia, North America, and northern Africa. Our results showed that body elemental content differs among B. terrestris castes. Young queens and workers had higher body nitrogen concentration than ovipositing queens and males, while castes did not differ significantly in their body carbon concentration. Furthermore, the carbon-to-nitrogen ratio was higher in ovipositing queens and males. We suggest that high body nitrogen concentration and low carbon-to-nitrogen ratio in young queens and workers may be related to their greater amount of flight muscles and flight activities than to their lower stress levels. To disentangle possible effects of stress in the agricultural landscape, further studies are needed to compare the elemental content of bumblebee bodies between natural habitats and areas of high-intensity agriculture.

Nano-CT imaging of larvae in the ant Pheidole hyatti reveals coordinated growth of a rudimentary organ necessary for soldier development.

The growth of imaginal discs in holometabolous insects is coordinated with larval growth to ensure the symmetrical and proportional development of the adult appendages. In ants, the differential growth of these discs generates distinct castes-the winged male and queen castes and the wingless worker caste. In the hyperdiverse ant genus Pheidole, the worker caste is composed of two morphologically distinct subcastes: small-headed minor workers and larger, big-headed, soldiers. Although these worker subcastes are completely wingless, soldier larvae develop rudimentary forewing discs that function in generating the disproportionate head-to-body scaling and size of soldiers. It remains unclear, however, how rudimentary forewing discs in soldier larvae are coordinated with other imaginal discs. Here we show, using quantitative nano-CT imaging and three-dimensional analyses, that the increase in the volume of the soldier rudimentary forewing discs is coordinated with larval size as well as with the increase in the volume of the leg and eye-antennal (head) discs. However, relative to larval size, we found that when the rudimentary forewing discs appear during the last larval instar, they are relatively smaller but increase in volume faster than that of the head (eye-antennal) and leg discs. These findings show that the rudimentary wing disc in soldier larvae has evolved novel patterns of inter-organ coordination as compared with other insects to generate the big-headed soldier caste in Pheidole. More generally, our study raises the possibility that novel patterns of inter-organ coordination are a general feature of rudimentary organs that acquire novel regulatory functions during development and evolution.

Impact of intraspecific variation on measurements of thermal tolerance in bumble bees.

Climate change is an important driver of bee decline despite the fact that many species might respond to climate change differently. One method to predict how a species will respond to climate change is to identify its thermal tolerance limits. However, differences in thermal tolerance might also occur among distant populations of the same species based on their local environment or even among castes of social insects. Here, we investigated intraspecific differences in thermal tolerance among subspecies of the large earth bumble bee, Bombus terrestris (Apidae). We determined the critical thermal minima and maxima (CTmin and CTmax, respectively) of workers and queens from three lab-reared B. terrestris subspecies (B. t. terrestris, B. t. audax, and B. t. canariensis) which originated from different thermal environments. Our results showed that caste has an influence on critical thermal minima, with queens being most cold-tolerant, but the values of critical thermal maxima were not correlated to caste or size. The thermal tolerance of workers did not differ among the subspecies. Although heat tolerance was similar in queens, B. t. canariensis queens (originating from the warmest environments) were the least cold tolerant. Overall, we showed that B. terrestris may be generally robust against climate warming, but that particular subspecies and/or populations may be more vulnerable to extreme temperature variability. Future research should focus on responses of B. terrestris populations to short, extreme thermal events.

The weird eusociality of polyembryonic parasites.

Some parasitoid wasps possess soldier castes during their parasitic larval stage, but are often neglected from our evolutionary theories explaining caste systems in animal societies. This is primarily due to the polyembryonic origin of their societies. However, recent discoveries of polyembryonic trematodes (i.e. flatworms) possessing soldier castes require us to reconsider this reasoning. I argue we can benefit from including these polyembryonic parasites in eusocial discussions, for polyembryony and parasitism are taxonomically vast and influence the evolution of social behaviours and caste systems in various circumstances. Despite their polyembryony, their social evolution can be explained by theories of eusociality designed for parent-offspring groups, which are the subjects of most social evolution research. Including polyembryonic parasites in these theories follows the trend of major evolutionary transitions theory expanding social evolution research into all levels of biological organization. In addition, these continued discoveries of caste systems in parasites suggest social evolution may be more relevant to parasitology than currently acknowledged.