Loss of Developmental Diapause as Prerequisite for Social Evolution in Bees

Diapause is a physiological arrest of development ahead of adverse environmental conditions and is a critical phase of the life cycle of many insects. In bees, diapause has been reported in species from all seven taxonomic families. However, they exhibit a variety of diapause strategies. These different strategies are of particular interest since shifts in the phase of the insect life cycle in which diapause occurs have been hypothesized to promote the evolution of sociality. Here we provide a comprehensive evaluation of this hypothesis with phylogenetic analysis and ancestral state reconstruction (ASR) of the ecological and evolutionary factors associated with diapause phase. We find that social lifestyle, latitude and voltinism are significant predictors of the life stage in which diapause occurs. ASR revealed that the most recent common ancestor of all bees likely exhibited developmental diapause and shifts to adult, reproductive, or no diapause have occurred in the ancestors of lineages in which social behaviour has evolved. These results provide fresh insight regarding the role of diapause as a prerequisite for the evolution of sociality in bees

Gregarious nesting of a digger wasp as a “selfish herd” response to a parasitic fly (Hymenoptera: Sphecidae; Diptera: Sacrophagidae)

Females of a digger wasp Crabro cribrellifer nested in a 234 nest aggregation covering 2x65 m in area. Variability in 3 edaphic factors (soil temperature, hardness, and per cent moisture) had no significant influence on the non-random spatial distribution of nests within the aggregation. A significant positive correlation between wasp nest density and the mean number of parasitic flies, Metopia campestris , present was shown (Fig. 1). Yet, based on 20 excavated nests, the probability of cell parasitism decreased as nest density increased (Fig. 2). This may give females nesting within a dense aggregation a selective advantage, and supports the “selfish herd” model developed by Hamilton (1971).Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/46872/1/265_2004_Article_BF00299384.pd

Gender Specific Brood Cells in the Solitary Bee Colletes halophilus (Hymenoptera; Colletidae)

We studied the reproductive behaviour of the solitary bee Colletes halophilus based on the variation in cell size, larval food amount and larval sex in relation to the sexual size dimorphism in this bee. Brood cells with female larvae are larger and contain more larval food than cells with males. Occasionally males are reared in female-sized cells. We conclude that a female C. halophilus in principal anticipates the sex of her offspring at the moment brood cell construction is started. Additionally a female is able to ‘change her mind’ about the sex of her offspring during a single brood cell cycle. We present a model that can predict the sex of the larvae in an early stage of development

Buzz pollination: studying bee vibrations on flowers

Approximately 6% of flowering plant species possess flowers with anthers that open through small pores or slits. Extracting pollen from this type of specialised flower is achieved most efficiently by vibrating the anthers, a behaviour that has evolved repeatedly among bees. Here I provide a brief overview of studying vibrations produced by bees and their effects on pollen release. I discuss how bee morphology and behaviour affect the mechanical properties of vibrations, and how floral traits may influence the transmission of those vibrations from the bee to the anther, thus mediating pollen release, and ultimately bee and plant fitness. I suggest that understanding the evolution of buzz pollination requires studying the biomechanics of bee vibrations and their transmission on flowers

Pollen Processing Behavior of Heliconius Butterflies: A Derived Grooming Behavior

Pollen feeding behaviors Heliconius and Laparus (Lepidoptera: Nymphalidae) represent a key innovation that has shaped other life history traits of these neotropical butterflies. Although all flower visiting Lepidoptera regularly come in contact with pollen, only Heliconius and Laparus butterflies actively collect pollen with the proboscis and subsequently take up nutrients from the pollen grains. This study focused on the behavior of pollen processing and compared the movement patterns with proboscis grooming behavior in various nymphalid butterflies using video analysis. The proboscis movements of pollen processing behavior consisted of a lengthy series of repeated coiling and uncoiling movements in a loosely coiled proboscis position combined with up and down movements and the release of saliva. The proboscis-grooming behavior was triggered by contamination of the proboscis in both pollen feeding and non-pollen feeding nymphalid butterflies. Proboscis grooming movements included interrupted series of coiling and uncoiling movements, characteristic sideways movements, proboscis lifting, and occasionally full extension of the proboscis. Discharge of saliva was more pronounced in pollen feeding species than in non-pollen feeding butterfly species. We conclude that the pollen processing behavior of Heliconius and Laparus is a modified proboscis grooming behavior that originally served to clean the proboscis after contamination with particles

Fossil Carder Bee's nest from the Hominin locality of Taung, South Africa

The Buxton-Norlim Limeworks southwest of Taung, South Africa, is renowned for the discovery of the first Australopithecus africanus fossil, the ‘Taung Child’. The hominin was recovered from a distinctive pink calcrete that contains an abundance of invertebrate ichnofauna belonging to the Coprinisphaera ichnofacies. Here we describe the first fossil bee’s nest, attributed to the ichnogenus Celliforma, from the Plio-Pleistocene of Africa. Petrographic examination of a cell lining revealed the preservation of an intricate organic matrix lined with the calcitic casts of numerous plant trichomes–a nesting behaviour unique to the modern-day carder bees (Anthidiini). The presence of Celliforma considered alongside several other recorded ichnofossils can be indicative of a dry, savannah environment, in agreement with recent work on the palaeoenvironment of Plio-Pleistocene southern Africa. Moreover, the occurrence of ground-nesting bees provides further evidence that the pink calcrete deposits are of pedogenic origin, rather than speleogenic origin as has previously been assumed. This study demonstrates the potential value of insect trace fossils as palaeoenvironmental indicators

Floral uniformity through evolutionary time in a species-rich tree lineage

Changes in floral morphology are expected across evolutionary time and are often promoted as important drivers in angiosperm diversification. Such a statement, however, is in contrast to empirical observations of species-rich lineages that show apparent conservative floral morphologies even under strong selective pressure to change from their environments. Here, we provide quantitative evidence for prolific speciation despite uniform floral morphology in a tropical species-rich tree lineage. We analyse floral disparity in the environmental and phylogenetic context of Myrcia (Myrtaceae), one of the most diverse and abundant tree genera in Neotropical biomes. Variation in floral morphology among Myrcia clades is exceptionally low, even among distantly related species. Discrete floral specialisations do occur, but these are few, present low phylogenetic signal, have no strong correlation with abiotic factors, and do not affect overall macroevolutionary dynamics in the lineage. Results show that floral form and function may be conserved over large evolutionary time scales even in environments full of opportunities for ecological interactions and niche specialisation. Species accumulation in diverse lineages with uniform flowers apparently does not result from shifts in pollination strategies, but from speciation mechanisms that involve other, nonfloral plant traits

The Antiquity and Evolutionary History of Social Behavior in Bees

A long-standing controversy in bee social evolution concerns whether highly eusocial behavior has evolved once or twice within the corbiculate Apidae. Corbiculate bees include the highly eusocial honey bees and stingless bees, the primitively eusocial bumble bees, and the predominantly solitary or communal orchid bees. Here we use a model-based approach to reconstruct the evolutionary history of eusociality and date the antiquity of eusocial behavior in apid bees, using a recent molecular phylogeny of the Apidae. We conclude that eusociality evolved once in the common ancestor of the corbiculate Apidae, advanced eusociality evolved independently in the honey and stingless bees, and that eusociality was lost in the orchid bees. Fossil-calibrated divergence time estimates reveal that eusociality first evolved at least 87 Mya (78 to 95 Mya) in the corbiculates, much earlier than in other groups of bees with less complex social behavior. These results provide a robust new evolutionary framework for studies of the organization and genetic basis of social behavior in honey bees and their relatives

At the brink of eusociality: transcriptomic correlates of worker behaviour in a small carpenter bee

Background: There is great interest in understanding the genomic underpinnings of social evolution, in particular, the evolution of eusociality (caste-containing societies with non-reproductives that care for siblings). Subsociality is a key precursor for the evolution of eusociality and characterized by prolonged parental care and parent-offspring interaction. Here, we provide the first transcriptomic data for the small carpenter bee, Ceratina calcarata. This species is of special interest because it is subsocial and in the same family as the highly eusocial honey bee, Apis mellifera. In addition, some C. calcarata females demonstrate alloparental care without reproduction, which provides a unique opportunity to study worker behaviour in a non-eusocial species. Results: We uncovered similar gene expression patterns related to maternal care and sibling care in different groups of females. This agrees with the maternal heterochrony hypothesis, specifically, that changes in timing of offspring care gene expression are related to worker behaviour in incipient insect societies. In addition, we also detected some similarity to caste-related gene expression patterns in highly eusocial honey bees, and uncovered large lifetime changes in gene expression that accompany shifts in reproductive and maternal care behaviour. Conclusions: For Ceratina calcarata, we found that transcript expression profiles were most similar between sibling care and maternal care females. The maternal care behaviour exhibited post-reproductively by Ceratina mothers is concordant in terms of transcript expression with the alloparental care exhibited by workers. In line with theoretical predictions, our data are consistent with the maternal heterochrony hypothesis for the evolutionary development of worker behaviour in subsocial bees