Evolution of Acquired Perfumes and Endogenous Lipid Secretions in Orchid Bees.

Male orchid bees are unique in the animal kingdom for making perfumes that function as sex pheromone. Males collect volatile chemicals from the environment in the neotropical forests, including floral and non-floral sources, creating complex but species-specific blends. Male orchid bees exhibit several adaptations to facilitate perfume collection and storage. When collecting volatile compounds, males apply lipid substances that they secrete from cephalic labial glands onto the fragrant substrate. These lipids help dissolve and retain the volatiles, similar to the process of 'enfleurage' in the traditional perfume industry. We investigated how the chemical composition of acquired perfume and labial gland secretions varied across the phylogeny of orchid bees, including 65 species in five genera from Central and South America. Perfumes showed rapid evolution as revealed by low overall phylogenetic signal, in agreement with the idea that perfume compounds diverge rapidly and substantially among closely related species due to their role in species recognition. A possible exception were perfumes in the genus Eulaema, clustering closely in chemospace, partly mediated by high proportions of carvone and trans-carvone oxide. Labial gland secretions, in contrast, showed a strong phylogenetic signal at the genus level, with secretions of Eufriesea and Exaerete dominated by fatty acids and Eulaema dominated by saturated acetates of chain lengths 12 to 16 C-atoms. Secretions of the majority of Euglossa were heavily dominated by one unsaturated long chain diacetate, (9Z)-Eicosen-1,20-diyldiacetate. However, we also identified few highly divergent species of Euglossa in four subclades (11 species) that appear to have secondarily replaced the diacetate with other compounds. In comparison with environment-derived perfumes, the evolution of labial gland secretion is much slower, likely constrained by the underlying biochemical pathways, but perhaps influenced by perfume-solvent chemical interactions.

Evidence for Reductions in Physical and Chemical Plant Defense Traits in Island Flora.

Reduced defense against large herbivores has been suggested to be part of the "island syndrome" in plants. However, empirical evidence for this pattern is mixed. In this paper, we present two studies that compare putative physical and chemical defense traits from plants on the California Channel Islands and nearby mainland based on sampling of both field and common garden plants. In the first study, we focus on five pairs of woody shrubs from three island and three mainland locations and find evidence for increased leaf area, decreased marginal leaf spines, and decreased concentrations of cyanogenic glycosides in island plants. We observed similar increases in leaf area and decreases in defense traits when comparing island and mainland genotypes grown together in botanic gardens, suggesting that trait differences are not solely driven by abiotic differences between island and mainland sites. In the second study, we conducted a common garden experiment with a perennial herb-Stachys bullata (Lamiaceae)-collected from two island and four mainland locations. Compared to their mainland relatives, island genotypes show highly reduced glandular trichomes and a nearly 100-fold reduction in mono- and sesquiterpene compounds from leaf surfaces. Island genotypes also had significantly higher specific leaf area, somewhat lower rates of gas exchange, and greater aboveground biomass than mainland genotypes across two years of study, potentially reflecting a broader shift in growth habit. Together, our results provide evidence for reduced expression of putative defense traits in island plants, though these results may reflect adaptation to both biotic (i.e., the historical absence of large herbivores) and climatic conditions on islands.

Chemical species recognition in an adaptive radiation of Hawaiian Tetragnatha spiders (Araneae: Tetragnathidae).

Studies of adaptive radiations have played a central role in our understanding of reproductive isolation. Yet the focus has been on human-biased visual and auditory signals, leaving gaps in our knowledge of other modalities. To date, studies on chemical signals in adaptive radiations have focused on systems with multimodal signalling, making it difficult to isolate the role chemicals play in reproductive isolation. In this study we examine the use of chemical signals in the species recognition and adaptive radiation of Hawaiian Tetragnatha spiders by focusing on entire communities of co-occurring species, and conducting behavioural assays in conjunction with chemical analysis of their silks using gas chromatography-mass spectrometry. Male spiders significantly preferred the silk extracts of conspecific mates over those of sympatric heterospecifics. The compounds found in the silk extracts, long chain alkyl methyl ethers, were remarkably species-specific in the combination and quantity. The differences in the profile were greatest between co-occurring species and between closely related sibling species. Lastly, there were significant differences in the chemical profile between two populations of a particular species. These findings provide key insights into the role chemical signals play in the attainment and maintenance of reproductive barriers between closely related co-occurring species.

The transcriptomic signature of adaptations associated with perfume collection in orchid bees.

Secondary sexual traits can convey information on mate quality with the signal honesty maintained by the costly nature of trait expression. Mating signals are also often underpinned by physiological, morphological, and behavioural adaptations, which may require the evolution of novelty, but the genetic basis in many cases is unknown. In orchid bees, males acquire chemical compounds from the environment that act as pheromone-like bouquets (perfumes) during courtship displays. This process could be costly, potentially due to the cognitive demands of learning and the physiological demands of collecting a mix of extrinsic chemical compounds that may require detoxification. Furthermore, a novel trait, a specialized perfume pouch in the hind leg, is required for compound storage. We studied gene expression in the brain, hind leg, and Malpighian tubules-a tissue involved in detoxification-to investigate changes in gene expression following perfume collection. We detected upregulation of genes enriched in functions related to transcription, odorant binding, and receptor activity in the Malpighian tubules. On the other hand, we did not find any evidence for learning processes following perfume collection, or gene expression changes in the hind leg, perhaps due to constitutive expression, or the age of the sampled bees. We did identify high expression of chemosensory proteins in the hind legs, which we suggest could play a role in perfume collection or storage, with further functional studies necessary to determine their binding properties and potential physiological importance. Los rasgos sexuales secundarios pueden servir como indicadores de calidad de la pareja, y en algunos casos la honestidad de la señal se mantiene por el costo de expresar el rasgo. A menudo las señales sexuales están respaldadas por adaptaciones fisiológicas, morfológicas y de comportamiento por lo tanto pueden requerir la evolución de nuevos rasgos, pero en muchos casos se desconoce la base genética. En las abejas de las orquídeas, los machos recolectan compuestos químicos del medio ambiente, los cuales actúan como feromonas (perfumes) durante el despliegue de cortejo. Este proceso podría ser costoso, posiblemente debido a las demandas cognitivas del aprendizaje y las demandas fisiológicas de recolectar una mezcla de compuestos químicos extrínsecos que pueden requerir desintoxicación. Además, se requiere la evolución de un contenedor para almacenar perfumes en la pata trasera. Para investigar los cambios en la expresión génica después de la recolección de perfume, estudiamos la expresión génica en el cerebro, la pata trasera y los túbulos de Malpighi (tejido involucrado en la desintoxicación). Encontramos varios genes regulados positivamente en los túbulos de Malpighi después de la recolección que están enriquecidos en factores de transcripción, proteínas de fijación de olores, y proteínas con actividad de receptor. Por otro lado, no encontramos ninguna evidencia de procesos de aprendizaje posteriores a la recolección de perfumes, o cambios en la expresión génica en la pata trasera, esto quizás debido a la expresión constitutiva o la edad de las abejas muestreadas. Además, identificamos una alta expresión de proteínas quimio-sensoriales en las patas traseras, que podría desempeñar un papel en la recolección o almacenamiento de perfumes. Más estudios funcionales son necesarios para determinar las propiedades de fijación de las proteínas y su potencial importancia fisiológica.

Macroevolution of floral scent chemistry across radiations of male euglossine bee-pollinated plantsMacroevolución de olores florales a través de radiaciones de plantas polinizadas por abejas euglosinas machosMacroevolução dos voláteis florais em radiações de plantas polinizadas por machos de abelhas Euglossini.

Floral volatiles play key roles as signaling agents that mediate interactions between plants and animals. Despite their importance, few studies have investigated broad patterns of volatile variation across groups of plants that share pollinators, particularly in a phylogenetic context. The "perfume flowers," Neotropical plant species exhibiting exclusive pollination by male euglossine bees in search of chemical rewards, present an intriguing system to investigate these patterns due to the unique function of their chemical phenotypes as both signaling agents and rewards. We leverage recently developed phylogenies and knowledge of biosynthesis, along with decades of chemical ecology research, to characterize axes of variation in the chemistry of perfume flowers, as well as understand their evolution at finer taxonomic scales. We detect pervasive chemical convergence, with many species across families exhibiting similar volatile phenotypes. Scent profiles of most species are dominated by compounds of either the phenylpropanoid or terpenoid biosynthesis pathways, while terpenoid compounds drive more subtle axes of variation. We find recapitulation of these patterns within two independent radiations of perfume flower orchids, in which we further detect evidence for the rapid evolution of divergent floral chemistries, consistent with the putative importance of scent in the process of adaptation and speciation.

Individual Variation in Male Pheromone Production in Xylocopa sonorina Correlates with size and Gland Color.

Sex pheromones are species-specific chemical signals that facilitate the location, identification, and selection of mating partners. These pheromones can vary between individuals, and act as signals of mate quality. Here, we investigate the variation of male pheromones in the mesosomal glands of the large carpenter bee Xylocopa sonorina, within a Northern California population. We tested the hypothesis that morphological traits are correlated with the observed variation in chemical blend composition of these bees. We also conducted behavioral assays to test whether these male pheromones act as long-range attractants to conspecifics. We found that larger males with darker mesosomal glands have a higher pheromone amount in their glands. Our analysis also suggests that this pheromone blend functions as a long-range attractant to both males and females. We show that both male body size and sexual maturation are important factors influencing pheromone abundance, and that this pheromone blend acts as a long-range attractant. We hypothesize that this recorded variation in male pheromone could be important for female choice.

Biogeography: The origin and spread of bee lineages.

Where and when bees originated and how they dispersed and diversified across ancient continents has remained ambiguous. A new study that combines phylogenetics with fossil data reconstructs the origin and diversification of bees across geological time and space.

Function of environment-derived male perfumes in orchid bees.

Perfume making in male orchid bees is a unique behavior that has given rise to an entire pollination syndrome in the neotropics.1,2 Male orchid bees concoct and store species-specific perfume mixtures in specialized hind-leg pockets3 using volatiles acquired from multiple environmental sources, including orchid flowers.4,5 However, the function and the ultimate causes of this behavior have remained elusive.2,6 Although previous observations suggested that male perfumes serve as chemical signals, the attractiveness for females has not be shown.7,8 Here, we demonstrate that the possession of perfume increases male mating success and paternity in Euglossa dilemma, a species of orchid bees recently naturalized in Florida. We supplemented males reared from trap-nests with perfume loads harvested from wild conspecifics. In dual-choice experiments, males supplemented with perfumes mated with more females, and sired more offspring, than untreated, equal-aged, control males. Although perfume supplementation had little effect on the intensity of male courtship display, it changed the dynamics of male-male interactions. Our results demonstrate that male-acquired perfumes are sexual signals that stimulate females for mating and suggest that sexual selection is key in shaping the evolution of perfume communication in orchid bees.

Seasonal stability and species specificity of environmentally acquired chemical mating signals in orchid bees.

Traits that mediate reproductive isolation between species, such as those involved in mate choice and/or recognition, are predicted to experience stabilizing selection towards the species mean. Male orchid bees collect chemical compounds from many sources, such as plants and fungi, which they use as a perfume signal (pheromone) during courtship display, and are suggested to contribute to reproductive isolation between species. Environmentally acquired signals are more prone to variation as source availability can vary through space and time. If orchid bee perfumes are important for reproductive isolation between species, we expect them to exhibit stable species-specific differences in time and space. Here, we describe phenotypic patterns of inter- and intraspecific variation in the male perfumes of three sympatric species of Euglossa orchid bees across an entire year, investigating both their seasonality and species specificity. Our analysis revealed considerable within-species variation in perfumes. However, species specificity was maintained consistently throughout the year, supporting the idea that these perfumes could play an important role in reproductive isolation and are experiencing stabilizing selection towards a species mean. Our analysis also identified strong correlations in the abundance of some compounds, possibly due to shared collection sources between species. Our study suggests that orchid bee perfumes are robust in the face of environmental changes in resource availability and thus can maintain reproductive isolation between species.

El aislamiento reproductivo entre especies es usualmente controlado por rasgos que juegan un papel importante en la selección y/o reconocimiento de pareja. Estos rasgos usualmente están bajo la presión de selección estabilizadora, lo que hace que se mantengan cerca al promedio de la especie. Los machos de las abejas de las orquídeas recolectan compuestos químicos de plantas y hongos y los usan como señales de feromonas durante el cortejo. Se piensa que las feromonas de las abejas de las orquídeas contribuyen al aislamiento reproductivo entre especies. Al ser obtenidas del ambiente, estas señales químicas (perfumes) son más propensas a la variación, ya que la disponibilidad de la fuente puede variar en el espacio y el tiempo. Si los perfumes contribuyen al aislamiento reproductivo, se esperaría que las diferencias entre especies sean estables tanto en el tiempo como en el espacio. Aquí, investigamos la estacionalidad y la especificidad a nivel de especie de los perfumes durante todo un año en tres especies simpátricas de abejas del género Euglossa. Nuestro análisis reveló variación considerable en los perfumes de cada especie. Sin embargo, la especificidad de cada especie se mantuvo durante todo el año, lo cual sugiere que los perfumes juegan un papel importante en el aislamiento reproductivo y que experimentan selección estabilizadora hacia la media de la especie. Nuestro análisis también identificó fuertes correlaciones en la abundancia de algunos compuestos, posiblemente debido a que diferentes especies comparten las mismas fuentes de recolección. Nuestro estudio sugiere que los perfumes de las abejas de las orquídeas son robustos frente a los cambios ambientales y la variación en la disponibilidad de recursos y por lo tanto pueden contribuir al aislamiento reproductivo.

Population genetics of a recent range expansion and subsequent loss of migration in monarch butterflies.

Range expansions-whether permanent or transient-strongly influence the distribution of genetic variation in space. Monarch butterflies are best known for long-distance seasonal migration within North America but are also established as nonmigratory populations around the world, including on Pacific Islands. Previous research has highlighted stepwise expansion across the Pacific, though questions remain about expansion timing and the population genetic consequences of migration loss. Here, we present reduced-representation sequencing data for 275 monarchs from North America (n = 85), 12 Pacific Islands (n = 136) and three locations in Australia (n = 54), with the goal of understanding (i) how the monarch's Pacific expansion has shaped patterns of population genetic variation and (ii) how loss of migration has influenced spatial patterns of differentiation. We find support for previously described stepwise dispersal across the Pacific and document an additional expansion from Hawaii into the Mariana Islands. Nonmigratory monarchs within the Mariana Islands show strong patterns of differentiation, despite their proximity; by contrast, migratory North American samples form a single genetically panmictic population across the continent. Estimates of Pacific establishment timing are highly uncertain (~100-1,000,000 years ago) but overlap with historical records that indicate a recent expansion. Our data support (i) a recent expansion across the Pacific whose timing overlaps with available historical records of establishment and (ii) a strong role for seasonal migration in determining patterns of spatial genetic variation. Our results are noteworthy because they demonstrate how the evolution of partial migration can drive population differentiation over contemporary timescales.

Experimental disruption of social structure reveals totipotency in the orchid bee, Euglossa dilemma.

Eusociality has evolved multiple times across the insect phylogeny. Social insects with greater levels of social complexity tend to exhibit specialized castes with low levels of individual phenotypic plasticity. In contrast, species with simple social groups may consist of totipotent individuals that transition among behavioral and reproductive states. However, recent work has shown that in simple social groups, there can still be constraint on individual plasticity, caused by differences in maternal nourishment or social interaction. It is not well understood how these constraints arise, ultimately leading to the evolution of nonreproductive workers. Some species of orchid bees form social groups of a dominant and-one to two subordinate helpers where all individuals are reproductive. Females can also disperse to start their own nest as a solitary foundress, which includes a nonreproductive phase characterized by ovary inactivation, not typically expressed by subordinates. Little is known about individual flexibility across these trajectories. Here, using the orchid bee Euglossa dilemma, we assess the plasticity of subordinate helpers, finding that they are capable of the same behavioral, physiological, transcriptomic, and chemical changes seen in foundresses. Our results suggest that the lack of nonreproductive workers in E. dilemma is not due to a lack of subordinate plasticity.

Evolution of Olfactory Receptors Tuned to Mustard Oils in Herbivorous Drosophilidae.

The diversity of herbivorous insects is attributed to their propensity to specialize on toxic plants. In an evolutionary twist, toxins betray the identity of their bearers when herbivores coopt them as cues for host-plant finding, but the evolutionary mechanisms underlying this phenomenon are poorly understood. We focused on Scaptomyza flava, an herbivorous drosophilid specialized on isothiocyanate (ITC)-producing (Brassicales) plants, and identified Or67b paralogs that were triplicated as mustard-specific herbivory evolved. Using in vivo heterologous systems for the expression of olfactory receptors, we found that S. flava Or67bs, but not the homologs from microbe-feeding relatives, responded selectively to ITCs, each paralog detecting different ITC subsets. Consistent with this, S. flava was attracted to ITCs, as was Drosophila melanogaster expressing S. flava Or67b3 in the homologous Or67b olfactory circuit. ITCs were likely coopted as olfactory attractants through gene duplication and functional specialization (neofunctionalization and subfunctionalization) in S. flava, a recently derived herbivore.

The Birth-and-Death Evolution of Cytochrome P450 Genes in Bees.

The birth-and-death model of multigene family evolution describes how gene families evolve and diversify through duplication and deletion. The cytochrome P450s are one of the most diverse and well-studied multigene families, involved in both physiological and xenobiotic functions. Extensive studies of insect P450 genes have demonstrated their role in insecticide resistance. Bees are thought to experience toxin exposure through their diet of nectar and pollen, as well as the resin-collecting behavior exhibited by some species. Here, we describe the repertoire of P450 genes in the orchid bee Euglossa dilemma. Male orchid bees form perfume bouquets used in courtship displays by collecting volatile compounds, resulting in exposure to compounds known to be toxic. In addition, we conducted phylogenetic and selection analyses across ten bee species encompassing three bee families. We find that social behavior and resin collection are not correlated with the repertoire of P450 present in a bee species. However, our analyses revealed that P450 clades can be classified as stable and unstable, and that genes involved in xenobiotic metabolism are more likely to belong to unstable clades. Furthermore, we find that unstable clades are under more dynamic evolutionary pressures and exhibit signals of adaptive evolution. This work highlights the complexity of multigene family evolution, revealing that multiple factors contribute to the diversification, stability, and dynamics of this gene family. Furthermore, we provide a resource for future detailed studies investigating the function of different P450s in economically important bee species.

Social Behavior, Ovary Size, and Population of Origin Influence Cuticular Hydrocarbons in the Orchid Bee Euglossa dilemma.

AbstractCuticular hydrocarbons (CHCs) are waxy compounds on the surface of insects that prevent desiccation and frequently serve as chemical signals mediating social and mating behaviors. Although their function in eusocial species has been heavily investigated, little is known about the evolution of CHC-based communication in species with simpler forms of social organization lacking specialized castes. Here we investigate factors shaping CHC variation in the orchid bee Euglossa dilemma, which forms casteless social groups of two to three individuals. We first assess geographic variation, examining CHC profiles of males and females from three populations. We also consider CHC variation in the sister species, Euglossa viridissima, which occurs sympatrically with one population of E. dilemma. Next, we consider variation associated with female behavioral phases, to test the hypothesis that CHCs reflect ovary size and social dominance. We uncover a striking CHC polymorphism in E. dilemma spanning populations. In addition, we identify a separate set of CHCs that correlate with ovary size, social dominance, and expression of genes associated with social behavior, suggesting that CHCs convey reproductive and social information in E. dilemma. Together, our results reveal complex patterns of variation in which a subset of CHCs reflect the social and reproductive status of nestmates.

Two centuries of monarch butterfly collections reveal contrasting effects of range expansion and migration loss on wing traits.

Migratory animals exhibit traits that allow them to exploit seasonally variable habitats. In environments where migration is no longer beneficial, such as oceanic islands, migration-association traits may be selected against or be under relaxed selection. Monarch butterflies are best known for their continent-scale migration in North America but have repeatedly become established as nonmigrants in the tropical Americas and on Atlantic and Pacific Islands. These replicated nonmigratory populations provide natural laboratories for understanding the rate of evolution of migration-associated traits. We measured >6,000 museum specimens of monarch butterflies collected from 1856 to the present as well as contemporary wild-caught monarchs from around the world. We determined 1) how wing morphology varies across the monarch's global range, 2) whether initial long-distance founders were particularly suited for migration, and 3) whether recently established nonmigrants show evidence for contemporary phenotypic evolution. We further reared >1,000 monarchs from six populations around the world under controlled conditions and measured migration-associated traits. Historical specimens show that 1) initial founders are well suited for long-distance movement and 2) loss of seasonal migration is associated with reductions in forewing size and elongation. Monarch butterflies raised in a common garden from four derived nonmigratory populations exhibit genetically based reductions in forewing size, consistent with a previous study. Our findings provide a compelling example of how migration-associated traits may be favored during the early stages of range expansion, and also the rate of reductions in those same traits upon loss of migration.

Selection and hybridization shaped the rapid spread of African honey bee ancestry in the Americas.

Recent biological invasions offer 'natural' laboratories to understand the genetics and ecology of adaptation, hybridization, and range limits. One of the most impressive and well-documented biological invasions of the 20th century began in 1957 when Apis mellifera scutellata honey bees swarmed out of managed experimental colonies in Brazil. This newly-imported subspecies, native to southern and eastern Africa, both hybridized with and out-competed previously-introduced European honey bee subspecies. Populations of scutellata-European hybrid honey bees rapidly expanded and spread across much of the Americas in less than 50 years. We use broad geographic sampling and whole genome sequencing of over 300 bees to map the distribution of scutellata ancestry where the northern and southern invasions have presently stalled, forming replicated hybrid zones with European bee populations in California and Argentina. California is much farther from Brazil, yet these hybrid zones occur at very similar latitudes, consistent with the invasion having reached a climate barrier. At these range limits, we observe genome-wide clines for scutellata ancestry, and parallel clines for wing length that span hundreds of kilometers, supporting a smooth transition from climates favoring scutellata-European hybrid bees to climates where they cannot survive winter. We find no large effect loci maintaining exceptionally steep ancestry transitions. Instead, we find most individual loci have concordant ancestry clines across South America, with a build-up of somewhat steeper clines in regions of the genome with low recombination rates, consistent with many loci of small effect contributing to climate-associated fitness trade-offs. Additionally, we find no substantial reductions in genetic diversity associated with rapid expansions nor complete dropout of scutellata ancestry at any individual loci on either continent, which suggests that the competitive fitness advantage of scutellata ancestry at lower latitudes has a polygenic basis and that scutellata-European hybrid bees maintained large population sizes during their invasion. To test for parallel selection across continents, we develop a null model that accounts for drift in ancestry frequencies during the rapid expansion. We identify several peaks within a larger genomic region where selection has pushed scutellata ancestry to high frequency hundreds of kilometers past the present cline centers in both North and South America and that may underlie high-fitness traits driving the invasion.

The evolution of sexual signaling is linked to odorant receptor tuning in perfume-collecting orchid bees.

Sexual signaling is an important reproductive barrier known to evolve early during the formation of new species, but the genetic mechanisms that facilitate the divergence of sexual signals remain elusive. Here we isolate a gene linked to the rapid evolution of a signaling trait in a pair of nascent neotropical orchid bee lineages, Euglossa dilemma and E. viridissima. Male orchid bees acquire chemical compounds from their environment to concoct species-specific perfumes to later expose during courtship. We find that the two lineages acquire chemically distinct perfumes and are reproductively isolated despite low levels of genome-wide differentiation. Remarkably, variation in perfume chemistry coincides with rapid divergence in few odorant receptor (OR) genes. Using functional assays, we demonstrate that the derived variant of Or41 in E. dilemma is specific towards its species-specific major perfume compound, whereas the ancestral variant in E. viridissima is broadly tuned to multiple odorants. Our results show that OR evolution likely played a role in the divergence of sexual communication in natural populations.

Host plant adaptation during contemporary range expansion in the monarch butterfly.

Herbivores that have recently expanded their host plant ranges provide opportunities to test hypotheses about the evolution of host plant specialization. Here, we take advantage of the contemporary global range expansion of the monarch butterfly (Danaus plexippus) and conduct a reciprocal rearing experiment involving monarch populations with divergent host plant assemblages. Specifically, we ask the following questions: (1) Do geographically disparate populations of monarch butterflies show evidence for local adaptation to their host plants? If so, what processes contribute to this pattern? (2) How is dietary breadth related to performance across multiple host species in monarch populations? (3) Does the coefficient of variation in performance vary across sympatric versus allopatric hosts? We find evidence for local adaptation in larval growth rate and survival based on sympatric/allopatric contrasts. Migratory North American monarchs, which have comparatively broad host breadth, have higher mean performance than derived nonmigratory populations across all host plant species. Monarchs reared on their sympatric host plants show lower coefficient of variation in performance than monarchs reared on allopatric hosts. We focus our discussion on possible mechanisms contributing to local adaptation to novel host plants and potential explanations for the reduction in performance that we observed in derived monarch populations.

Sociality emerges from solitary behaviours and reproductive plasticity in the orchid bee Euglossa dilemma.

The evolution of eusociality and sterile worker castes represents a major transition in the history of life. Despite this, little is known about the mechanisms involved in the initial transition from solitary to social behaviour. It has been hypothesized that plasticity from ancestral solitary life cycles was coopted to create queen and worker castes in insect societies. Here, we tested this hypothesis by examining gene expression involved in the transition from solitary to social behaviour in the orchid bee Euglossa dilemma. To this end, we conducted observations that allowed us to classify bees into four distinct categories of solitary and social behaviour. Then, by sequencing brain and ovary transcriptomes from these behavioural phases, we identified gene expression changes overlapping with socially associated genes across multiple eusocial lineages. We find that genes involved in solitary E. dilemma ovarian plasticity overlap extensively with genes showing differential expression between fertile and sterile workers-or between queens and workers in other eusocial bees. We also find evidence that sociality in E. dilemma reflects gene expression patterns involved in solitary foraging and non-foraging nest care behaviours. Our results provide strong support for the hypothesis that eusociality emerges from plasticity found across solitary life cycles.

Sexual dimorphism in visual and olfactory brain centers in the perfume-collecting orchid bee Euglossa dilemma (Hymenoptera, Apidae).

Insect mating behavior is controlled by a diverse array of sex-specific traits and strategies that evolved to maximize mating success. Orchid bees exhibit a unique suite of perfume-mediated mating behaviors. Male bees collect volatile compounds from their environment to concoct species-specific perfume mixtures that are presumably used to attract conspecific females. Despite a growing understanding of the ecology and evolution of chemical signaling in orchid bees, many aspects of the functional adaptations involved, in particular regarding sensory systems, remain unknown. Here we investigated male and female brain morphology in the common orchid bee Euglossa dilemma Bembé & Eltz. Males exhibited increased relative volumes of the Medulla, a visual brain region, which correlated with larger compound eye size (area). While the overall volume of olfactory brain regions was similar between sexes, the antennal lobes exhibited several sex-specific structures including one male-specific macroglomerulus. These findings reveal sexual dimorphism in both the visual and the olfactory system of orchid bees. It highlights the tendency of an increased investment in the male visual system similar to that observed in other bee lineages, and suggests that visual input may play a more important role in orchid bee male mating behavior than previously thought. Furthermore, our results suggest that the evolution of perfume communication in orchid bees did not involve drastic changes in olfactory brain morphology compared to other bee lineages.