Trade‐offs in the evolution of bumblebee colony and body size: a comparative analysis

Trade‐offs between life‐history traits – such as fecundity and survival – have been demonstrated in several studies. In eusocial insects, the number of organisms and their body sizes can affect the fitness of the colony. Large‐than‐average body sizes as well as more individuals can improve a colony's thermoregulation, foraging efficiency, and fecundity. However, in bumblebees, large colonies and large body sizes depend largely on high temperatures and a large amount of food resources. Bumblebee taxa can be found in temperate and tropical regions of the world and differ markedly in their colony sizes and body sizes. Variation in colony size and body size may be explained by the costs and benefits associated with the evolutionary history of each species in a particular environment. In this study, we explored the effect of temperature and precipitation (the latter was used as an indirect indicator of food availability) on the colony and body size of twenty‐one bumblebee taxa. A comparative analysis controlling for phylogenetic effects as well as for the body size of queens, workers, and males in bumblebee taxa from temperate and tropical regions indicated that both temperature and precipitation affect colony and body size. We found a negative association between colony size and the rainiest trimester, and a positive association between the colony size and the warmest month of the year. In addition, male bumblebees tend to evolve larger body sizes in places where the rain occurs mostly in the summer and the overall temperature is warmer. Moreover, we found a negative relationship between colony size and body sizes of queens, workers, and males, suggesting potential trade‐offs in the evolution of bumblebee colony and body size.     

The combined effect of host and food availability on optimized parasitoid life‐history traits based on a three‐dimensional trade‐off surface

The reproductive success of many insects is considered to be limited by two main factors: the availability of mature eggs to lay (termed egg limitation) and the time to locate suitable hosts (termed time limitation). High host density in the environment is likely to enhance oviposition opportunities, thereby selecting for higher investment in egg supply. In contrast, a shortage of food (e.g. sugar sources) is likely to increase the risk of time limitation, thereby selecting for higher allocation to initial energy reserves. To our knowledge, the combined effect of host and food availability on these optimal life‐history allocations has never been investigated. We thus modelled their simultaneous effects on a three‐dimensional trade‐off between initial investment in energy reserves, egg number and egg size, while focusing on insect parasitoids. The model was based on Monte Carlo simulations coupled with genetic algorithms, in order to identify the optimal life‐history traits of a single simulated parasitoid female in an environment in which both hosts and food are present in varying densities. Our results reproduced the simple predictions described above. However, some novel predictions were also obtained, especially when specific interactions between the different factors were examined and their effects on the three‐dimensional life‐history surface were considered. The work sheds light on long‐lasting debates regarding the relative importance of time versus egg limitation in determining insect life‐history traits and highlights the complexity of life‐history evolution, where several environmental factors act simultaneously on multiple traits.     

Temperature‐mediated trade‐offs and changes in life‐history integration in two slipper limpets (Gastropoda: Calyptraeidae) with planktotrophic development

Intraspecific variation in egg size and hatching size, and the genetic and environmental trade‐offs that contribute to variation, are the basis of the evolution of life histories. The present study examined both univariate and multivariate temperature‐mediated plasticity of life‐history traits, as well as temperature‐mediated trade‐offs in egg size and clutch size, in two planktotrophic species of marine slipper limpets, Crepidula. Previous work with two species of Crepidula with large eggs and lecithotrophic development has shown a significant effect of temperature on egg size and hatching size. To further examine the effect of temperature on egg size in Crepidula, the effects of temperature on egg size and hatching size, as well as the possible trade‐offs with other the life‐history features, were examined for two planktotrophic species: Crepidula incurva and Crepidula cf. marginalis. Field‐collected juveniles were raised at 23 or 28 °C and egg size, hatching size, capsules/brood, eggs/capsule, time to hatch, interbrood interval, and final body weight were recorded. Consistent with results for the lecithotrophic Crepidula, egg size and hatching size decreased with temperature in the planktotrophic species. The affects of maternal identity and individual brood account for more than half of the intraspecific variation in egg size and hatching size. Temperature also showed a significant effect on reproductive rate, with time to hatch and interbrood interval both decreasing with increasing temperature. However, temperature had contrasting effects on the number of offspring. Crepidula cf. marginalis has significantly more eggs/capsule and therefore more eggs per brood at 28 °C compared to 23 °C, although capsules/brood did not vary with temperature. Crepidula incurva, on the other hand, produced significantly more capsules/brood and more eggs per brood at the lower temperature, whereas the number of eggs/capsule did not vary with temperature. The phenotypic variance–covariance matrix of life‐history variables showed a greater response to temperature in C. incurva than in C. cf. marginalis, and temperature induced trade‐offs between offspring size and number differ between the species. These differences suggest that temperature changes as a result of seasonal upwelling along the coast of Panama will effect the reproduction and evolution of life histories of these two co‐occurring species differently. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Size‐induced phenotypic reaction norms in a parasitoid wasp: an examination of life‐history and behavioural traits

The amount of resources available during development often affects body size, causing phenotypic variation in life‐history traits and reproductive behaviours. However, past studies have seldom examined the reaction norms of both life‐history and behavioural traits versus body size. We measured the phenotypic plasticity of several life‐history (age‐specific egg load, egg size, longevity) and behavioural (oviposition rate, host marking rate, walking speed) traits of the egg parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae) in response to body size variation. We predicted that life‐history traits would show more evidence of size compensation than behavioural traits, resulting in fewer positively‐sloped size versus trait reaction norms among the former. As predicted by life‐history models, smaller wasps appear to shift resource allocation towards early‐life reproduction, having a similar egg load to large individuals 9 days after emergence. Surprisingly, longevity was unaffected by body size. However, egg size, the number of offspring produced during oviposition bouts, and the rate of subsequent egg synthesis were greater for larger individuals. In addition, as predicted, the reaction norms of behavioural traits versus body size were all positively sloped. Thus, despite possible adaptive compensatory plasticity of life‐history traits by small individuals, behavioural constraints directly related to body size would contribute to maintaining a positive size–fitness relationship.     

South temperate birds have higher apparent adult survival than tropical birds in Africa

Life history theory predicts an inverse relationship between annual adult survival and fecundity. Globally, clutch size shows a latitudinal gradient among birds, with south temperate species laying smaller clutches than north temperate species, but larger clutches than tropical species. Tropical birds often have higher adult survival than north temperate birds associated with their smaller clutches. However, the prediction that tropical birds should also have higher adult survival than south temperate birds because of smaller clutch sizes remains largely untested. We measured clutch size and apparent annual breeding adult survival for 17 south temperate African species to test two main predictions. First, we found strong support for a predicted inverse relationship between adult survival and clutch size among the south temperate species, consistent with life‐history theory. Second, we compared our clutch size and survival estimates with published estimates for congeneric tropical African species to test the prediction of larger clutch size and lower adult survival among south temperate than related tropical species. We found that south‐temperate species laid larger clutches, as predicted, but had higher, rather than lower, apparent adult survival than related tropical species. The latter result may be an artefact of different approaches to measuring survival, but the results suggest that adult survival is generally high in the south temperate region and raises questions about the importance of the cost of reproduction to adult survival.     

How do host plant use and seasonal life cycle relate to insect body size: A case study on European geometrid moths (Lepidoptera: Geometridae)

We used European geometrid moths (>630 species) as a model group to investigate how life history traits linked to larval host plant use (i.e., diet breadth and host-plant growth form) and seasonal life cycle (i.e., voltinism, overwintering stage and caterpillar phenology) are related to adult body size in holometabolous insect herbivores. To do so, we applied phylogenetic comparative methods to account for shared evolutionary history among herbivore species. We further categorized larval diet breadth based on the phylogenetic structure of utilized host plant genera. Our results indicate that species associated with woody plants are, on average, larger than herb feeders and increase in size with increasing diet breadth. Obligatorily univoltine species are larger than multivoltine species, and attain larger sizes when their larvae occur exclusively in the early season. Furthermore, the adult body size is significantly smaller in species that overwinter in the pupal stage compared to those that overwinter as eggs or caterpillars. In summary, our results indicate that the ecological niche of holometabolous insect herbivores is strongly interrelated with body size at maturity.     

Costs of reproduction can explain the correlated evolution of semelparity and egg size: theory and a test with salmon

Species’ life history traits, including maturation age, number of reproductive bouts, offspring size and number, reflect adaptations to diverse biotic and abiotic selection pressures. A striking example of divergent life histories is the evolution of either iteroparity (breeding multiple times) or semelparity (breed once and die). We analysed published data on salmonid fishes and found that semelparous species produce larger eggs, that egg size and number increase with salmonid body size among populations and species and that migratory behaviour and parity interact. We developed three hypotheses that might explain the patterns in our data and evaluated them in a stage‐structured modelling framework accounting for different growth and survival scenarios. Our models predict the observation of small eggs in iteroparous species when egg size is costly to maternal survival or egg number is constrained. By exploring trait co‐variation in salmonids, we generate new hypotheses for the evolution of trade‐offs among life history traits.     

Selection and constraints on offspring size‐number trade‐offs in sand lizards (Lacerta agilis)

The trade‐off between offspring size and number is a central component of life‐history theory, postulating that larger investment into offspring size inevitably decreases offspring number. This trade‐off is generally discussed in terms of genetic, physiological or morphological constraints; however, as among‐individual differences can mask individual trade‐offs, the underlying mechanisms may be difficult to reveal. In this study, we use multivariate analyses to investigate whether there is a trade‐off between offspring size and number in a population of sand lizards by separating among‐ and within‐individual patterns using a 15‐year data set collected in the wild. We also explore the ecological and evolutionary causes and consequences of this trade‐off by investigating how a female's resource (condition)‐ vs. age‐related size (snout‐vent length) influences her investment into offspring size vs. number (OSN), whether these traits are heritable and under selection and whether the OSN trade‐off has a genetic component. We found a negative correlation between offspring size and number within individual females and physical constraints (size of body cavity) appear to limit the number of eggs that a female can produce. This suggests that the OSN trade‐off occurs due to resource constraints as a female continues to grow throughout life and, thus, produces larger clutches. In contrast to the assumptions of classic OSN theory, we did not detect selection on offspring size; however, there was directional selection for larger clutch sizes. The repeatabilities of both offspring size and number were low and we did not detect any additive genetic variance in either trait. This could be due to strong selection (past or current) on these life‐history traits, or to insufficient statistical power to detect significant additive genetic effects. Overall, the findings of this study are an important illustration of how analyses of within‐individual patterns can reveal trade‐offs and their underlying causes, with potential evolutionary and ecological consequences that are otherwise hidden by among‐individual variation.     

The effect of multi‐species host density on superparasitism and sex ratio in a gregarious parasitoid

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Elevational trends in life histories: revising the pace‐of‐life framework

Life‐history traits in birds, such as lifespan, age at maturity, and rate of reproduction, vary across environments and in combinations imposed by trade‐offs and limitations of physiological mechanisms. A plethora of studies have described the diversity of traits and hypothesized selection pressures shaping components of the survival–reproduction trade‐off. Life‐history variation appears to fall along a slow–fast continuum, with slow pace characterized by higher investment in survival over reproduction and fast pace characterized by higher investment in reproduction over survival. The Pace‐of‐Life Syndrome (POLS) is a framework to describe the slow–fast axis of variation in life‐history traits and physiological traits. The POLS corresponds to latitudinal gradients, with tropical birds exhibiting a slow pace of life. We examined four possible ways that the traits of high‐elevation birds might correspond to the POLS continuum: (i) rapid pace, (ii) tropical slow pace, (iii) novel elevational pace, or (iv) constrained pace. Recent studies reveal that birds breeding at high elevations in temperate zones exhibit a combination of traits creating a unique elevational pace of life with a central trade‐off similar to a slow pace but physiological trade‐offs more similar to a fast pace. A paucity of studies prevents consideration of the possibility of a constrained pace of life. We propose extending the POLS framework to include trait variation of elevational clines to help to investigate complexity in global geographic patterns.     

Egg‐size plasticity in Apis mellifera: Honey bee queens alter egg size in response to both genetic and environmental factors

Social evolution has led to distinct life‐history patterns in social insects, but many colony‐level and individual traits, such as egg size, are not sufficiently understood. Thus, a series of experiments was performed to study the effects of genotypes, colony size and colony nutrition on variation in egg size produced by honey bee (Apis mellifera) queens. Queens from different genetic stocks produced significantly different egg sizes under similar environmental conditions, indicating standing genetic variation for egg size that allows for adaptive evolutionary change. Further investigations revealed that eggs produced by queens in large colonies were consistently smaller than eggs produced in small colonies, and queens dynamically adjusted egg size in relation to colony size. Similarly, queens increased egg size in response to food deprivation. These results could not be solely explained by different numbers of eggs produced in the different circumstances but instead seem to reflect an active adjustment of resource allocation by the queen in response to colony conditions. As a result, larger eggs experienced higher subsequent survival than smaller eggs, suggesting that honey bee queens might increase egg size under unfavourable conditions to enhance brood survival and to minimize costly brood care of eggs that fail to successfully develop, and thus conserve energy at the colony level. The extensive plasticity and genetic variation of egg size in honey bees has important implications for understanding life‐history evolution in a social context and implies this neglected life‐history stage in honey bees may have trans‐generational effects.     

Contrasting latitudinal patterns of life‐history divergence in two genera of new world thrushes (Turdinae)

Several long‐standing hypotheses have been proposed to explain latitudinal patterns of life‐history strategies. Here, we test predictions of four such hypotheses (seasonality, food limitation, nest predation and adult survival probability) by examining life‐history traits and age‐specific mortality rates of several species of thrushes (Turdinae) based on field studies at temperate and tropical sites and data gathered from the literature. Thrushes in the genus Catharus showed the typical pattern of slower life‐history strategies in the tropics while co‐occuring Turdus thrushes differed much less across latitudes. Seasonality is a broadly accepted hypothesis for latitudinal patterns, but the lack of concordance in latitudinal patterns between co‐existing genera that experience the same seasonal patterns suggests seasonality cannot fully explain latitudinal trait variation in thrushes. Nest‐predation also could not explain patterns based on our field data and literature data for these two genera. Total feeding rates were similar, and per‐nestling feeding rates were higher at tropical latitudes in both genera, suggesting food limitation does not explain trait differences in thrushes. Latitudinal patterns of life histories in these two genera were closely associated with adult survival probability. Thus, our data suggest that environmental influences on adult survival probability may play a particularly strong role in shaping latitudinal patterns of life‐history traits.     

Life‐history traits and climatic responsiveness in noctuid moths

Emergence phenology has been shown to advance considerably in the past decades in many lepidopterans. Noctuid moths (Noctuidae) constitute a species‐rich family of lepidopterans with a large diversity of life history traits presumably driving climatic responsiveness. In our study we aim to assess the role of life‐history and ecological traits in climatic responsiveness of noctuid moths, whilst controlling for phylogenetic dependence. We used a long‐term dataset of European noctuid moths collected from a light‐trap in northeastern Hungary. As the study site is located at the intersection of several biogeographical zones harbouring a large number of noctuid moth species, our dataset provides a unique possibility to investigate the moths’ climatic sensitivity. To estimate the role of life‐history traits and ecological factors in driving lepidopterans’ response to climatic trends, we employed three proxies related to the species’ ecology (habitat affinity, food plant specialization and food type) and two robust types of life‐history traits (migration strategy and hibernation form). The degree of temporal shifts of various measures of emergence phenology was related to hibernation stage, food type and migration strategy. Large‐scale phylogenetic relatedness exerted little constraint in all models fitted on each measure of phenology. Our results imply that noctuid moths overwintering as adults exhibited greater degrees of phenological shifts than species hibernating as larvae or pupae. It implies that moths hibernating as adults are forced to suspend activity in our climate and the prolongation of autumn activity might be the result of increased plasticity in flight periods.     

Sexual dimorphism explains residual variance around the survival‐reproduction tradeoff in lizards: Implications for sexual conflict over life‐history evolution*

The tradeoff between survival and reproduction is a central feature of life‐history variation, but few studies have sought to explain why females of some species exhibit relatively lower survival than expected for a given level of reproductive effort (RE). Intralocus sexual conflict theory proposes that sex differences in selection on survival and RE may, by virtue of shared genes underlying these components of fitness, prevent females from optimizing this life‐history tradeoff. To test this hypothesis, we used a phylogenetically based comparative analysis of published estimates for mean annual survival and RE from females of 82 lizard species to (1) characterize the tradeoff between survival and reproduction and (2) test whether variation around this tradeoff is explained by sexual size dimorphism (SSD), a potential proxy for sexual conflict over life‐history traits. Across species, we found a strong negative correlation between mean annual survival and RE, confirming this classic life‐history tradeoff. Although residual variance around this tradeoff is unrelated to the absolute magnitude of SSD, it is strongly related to the direction of SSD. Specifically, we found that females have lower survival than expected for a given level of RE in female‐larger species, whereas they have higher survival than expected in male‐larger species. Given that female‐larger SSD is thought to reflect selection for increased fecundity, our results suggest that intralocus sexual conflict may be particularly likely to constrain female life‐history evolution in situations where increased RE is favored, but the phenotypes that facilitate this increase (e.g., body size) are constrained by antagonistic selection on males.     

Maternal and environmental influences on egg size and juvenile life‐history traits in Pacific salmon

Life‐history traits such as fecundity and offspring size are shaped by investment trade‐offs faced by mothers and mediated by environmental conditions. We use a 21‐year time series for three populations of wild sockeye salmon (Oncorhynchus nerka) to test predictions for such trade‐offs and responses to conditions faced by females during migration, and offspring during incubation. In years when their 1100 km upstream migration was challenged by high water discharges, females that reached spawning streams had invested less in gonads by producing smaller but not fewer eggs. These smaller eggs produced lighter juveniles, and this effect was further amplified in years when the incubation water was warm. This latter result suggests that there should be selection for larger eggs to compensate in populations that consistently experience warm incubation temperatures. A comparison among 16 populations, with matching migration and rearing environments but different incubation environments (i.e., separate spawning streams), confirmed this prediction; smaller females produced larger eggs for their size in warmer creeks. Taken together, these results reveal how maternal phenotype and environmental conditions can shape patterns of reproductive investment and consequently juvenile fitness‐related traits within and among populations.     

Age‐ and sex‐related differences in the spatial ecology of a dichromatic tropical python (Morelia viridis)

Abstract: Despite outnumbering their temperate counterparts, tropical snake species have been poorly studied. Yet, the few tropical species that have been studied show a variety of behavioural traits beyond those described in temperate species. Here we reveal both age and sexual differences in the movements of tropical green pythons (Morelia viridis: Pythonidae). We radio‐tracked 27 individuals (17 females and 10 males) for up to 18 months, locating individuals during both the day and night. The home range size for adult females (mean ± SE of 6.21 ± 1.85 ha) was correlated with snout–vent length. Neither adult males nor juveniles had a stable home range. Adult females had stable home ranges that overlapped considerably with those of other females and yellow individuals. Multiple radio‐tracked adult males passed through the territory of radio‐tracked adult females during the study. Females of all sizes were more likely to change position each day than males. There were no differences between the sexes or size categories in the distances moved in most months, although the variation in movement distances was higher in the dry season than during the wet season. In the wet season (January–March) movement distances increased and these were size‐ and sex‐related. This increased activity may be associated with mate searching. Males of M. viridis may maximize their rate of encountering mature females by roaming rather than maintaining a home range. Juvenile green pythons moved distances equal to adult snakes in most months despite their comparatively small size.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Interspecific associations between circulating antioxidant levels and life-history variation in birds

Antioxidants play an important role in protecting tissues against aging-associated oxidative damage and are thus prime candidates for relating physiological mechanisms to variation in life histories. We measured total antioxidant capacity, antioxidant response to stress, and levels of uric acid, vitamin E, and four carotenoids in 95 avian species, mostly passerines from Michigan or Panama. We compared antioxidant measures to seven variables related to life histories (clutch size, survival rate, incubation period, nestling period, basal metabolic rate, body mass, and whether the species lived in a tropical or temperate climate). Life-history-related traits varied over at least three statistically independent axes. Higher antioxidant levels were generally characteristic of more rapid development, lower survival rate, smaller body size, larger clutch size, and higher mass-adjusted metabolic rate, but the relationships of particular antioxidants with individual life-history traits showed considerable complexity. Antioxidant-life history associations differed between tropical and temperate species and varied with respect to taxonomic sampling. Vitamin E showed few relationships with life-history traits. Overall, our results partly support the hypothesis that antioxidant levels evolve to mirror free radical production. Clearly, however, the complex patterns of physiological diversification observed here result from the interplay of many factors, likely including not just investment in somatic maintenance but also phylogenetic constraint, diet, and other aspects of ecology.     

A comparative perspective on longevity: the effect of body size dominates over ecology in moths

Both physiologically and ecologically based explanations have been proposed to account for among‐species differences in lifespan, but they remain poorly tested. Phylogenetically explicit comparative analyses are still scarce and those that exist are biased towards homoeothermic vertebrates. Insect studies can significantly contribute as lifespan can feasibly be measured in a high number of species, and the selective forces that have shaped it may differ largely between species and from those acting on larger animals. We recorded adult lifespan in 98 species of geometrid moths. Phylogenetic comparative analyses were applied to study variation in species‐specific values of lifespan and to reveal its ecological and life‐history correlates. Among‐species and between‐gender differences in lifespan were found to be notably limited; there was also no evidence of phylogenetic signal in this trait. Larger moth species were found to live longer, with this result supporting a physiological rather than ecological explanation of this relationship. Species‐specific lifespan values could not be explained by traits such as reproductive season and larval diet breadth, strengthening the evidence for the dominance of physiological determinants of longevity over ecological ones.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.