The quantitative genetics of two life history trade-offs in the yellow dung fly in abundant and limited food environments

The trade-offs between body size and development time and between egg size and egg number (clutch size) are central to life history theory, but evidence for them, particularly in terms of genetic correlations, is equivocal. For the yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae), we investigated variation in phenotypic and genetic variances and covariances, i.e. heritabilities and genetic correlations, of these life history traits (plus diapause) in benign and stressful larval field or adult laboratory food environments. We found both trade-offs to be weak, as evidenced by low phenotypic and genetic correlations, but stronger in the food limited environments. Broad sense heritabilities were generally significant for all traits considered, whereas the narrow sense heritabilities for egg and clutch size were nil. With regard to the question of how environmental stress affects heritabilities, we found a whole range of responses within one single species depending on the traits considered. All three possible patterns occurred, i.e. increased h2 due to increased VG or decreased $$V_{P^{\prime}}$$ decreased h2 due to increased $$V_{P^{\prime}}$$ and no change in h2 due to increased VG and VP. These can be explained by the particular ecological circumstances yellow dung flies face in their natural environment. Nevertheless, the majority of patterns was consistent with the idea that stressful conditions amplify phenotypic differences between genotypes. Such variable responses of traits even within one organism underscores the complexity of this issue and may well explain the multiple patterns found in various organism

Bergmann and Converse Bergmann Latitudinal Clines in Arthropods: Two Ends of a Continuum?

Two seemingly opposite evolutionary patterns of clinal variation in body size and associated life history traits exist in nature. According to Bergmann's rule, body size increases with latitude, a temperature effect. According to the converse Bergmann rule, body size decreases with latitude, a season length effect. A third pattern causally related to the latter is countergradient variation, whereby populations of a given species compensate seasonal limitations at higher latitudes by evolving faster growth and larger body sizes compared to their low latitude conspecifics. We discuss these patterns and argue that they are not mutually exclusive because they are driven by different environmental causes and proximate mechanisms; they therefore can act in conjunction, resulting in any intermediate pattern. Alternatively, Bergmann and converse Bergmann clines can be interpreted as over- and undercompensating countergradient variation, respectively. We illustrate this with data for the wide-spread yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), which in Europe shows a Bergmann cline for size and a converse Bergmann cline (i.e., countergradient variation) for development time. A literature review of the available evidence on arthropod latitudinal clines further shows a patterned continuum of responses. Converse Bergmann clines due to end-of-season time limitations are more common in larger species with longer development times. Our study thus provides a synthesis to the controversy about the importance of Bergmann's rule and the converse Bergmann rule in natur

Environmental influences on the gametic investment of yellow dung fly males

The energetic investment per spermatozoon and in spermatogenesis is central to a male's reproductive strategy. Relatively little, however, is known about environmental influences on variation in male allocation decisions and associated trade-offs. Plasticity in sperm length and testis size in response to variable food and temperature conditions either before or after adult eclosion was investigated in Scathophaga stercoraria, a classic model organism for sperm competition. Both measures showed interesting and clear environmental effects and also a heritable component. Testis length, and thus presumably sperm production, showed a hypoallometric (b < 1), but non-linear increase with body size, indicating that the allometric relationship changed with size. Like body size, testis length decreased with increasing developmental temperatures, but also showed a complex cubic relationship with adult temperatures. In contrast, sperm length increased or showed a negative quadratic relationship with increasing temperatures. The increase of within-male variation in sperm length with increasing developmental temperature and decreasing adult food indicates that some of our treatments were stressful. Nevertheless, there was no evidence of a trade-off between testis size and sperm length. The missing effect of adult or larval food availability on testis and sperm length, despite strong effects of larval food on body size, suggests that investment into reproduction is less sensitive to food restriction than investment into growt

Bergmann and converse bergmann latitudinal clines in arthropods: two ends of a continuum?

Two seemingly opposite evolutionary patterns of clinal variation in body size and associated life history traits exist in nature. According to Bergmann's rule, body size increases with latitude, a temperature effect. According to the converse Bergmann rule, body size decreases with latitude, a season length effect. A third pattern causally related to the latter is countergradient variation, whereby populations of a given species compensate seasonal limitations at higher latitudes by evolving faster growth and larger body sizes compared to their low latitude conspecifics. We discuss these patterns and argue that they are not mutually exclusive because they are driven by different environmental causes and proximate mechanisms; they therefore can act in conjunction, resulting in any intermediate pattern. Alternatively, Bergmann and converse Bergmann clines can be interpreted as over- and undercompensating countergradient variation, respectively. We illustrate this with data for the wide-spread yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), which in Europe shows a Bergmann cline for size and a converse Bergmann cline (i.e., countergradient variation) for development time. A literature review of the available evidence on arthropod latitudinal clines further shows a patterned continuum of responses. Converse Bergmann clines due to end-of-season time limitations are more common in larger species with longer development times. Our study thus provides a synthesis to the controversy about the importance of Bergmann's rule and the converse Bergmann rule in natur

Sex-biased parental care and sexual size dimorphism in a provisioning arthropod

The diverse selection pressures driving the evolution of sexual size dimorphism (SSD) have long been debated. While the balance between fecundity selection and sexual selection has received much attention, explanations based on sex-specific ecology have proven harder to test. In ectotherms, females are typically larger than males, and this is frequently thought to be because size constrains female fecundity more than it constrains male mating success. However, SSD could additionally reflect maternal care strategies. Under this hypothesis, females are relatively larger where reproduction requires greater maximum maternal effort – for example where mothers transport heavy provisions to nests. To test this hypothesis we focussed on digger wasps (Hymenoptera: Ammophilini), a relatively homogeneous group in which only females provision offspring. In some species, a single large prey item, up to 10 times the mother’s weight, must be carried to each burrow on foot; other species provide many small prey, each flown individually to the nest. We found more pronounced female-biased SSD in species where females carry single, heavy prey. More generally, SSD was negatively correlated with numbers of prey provided per offspring. Females provisioning multiple small items had longer wings and thoraxes, probably because smaller prey are carried in flight. Despite much theorising, few empirical studies have tested how sex-biased parental care can affect SSD. Our study reveals that such costs can be associated with the evolution of dimorphism, and this should be investigated in other clades where parental care costs differ between sexes and species

The impacts of environmental warming on Odonata: a review

Climate change brings with it unprecedented rates of increase in environmental temperature, which will have major consequences for the earth's flora and fauna. The Odonata represent a taxon that has many strong links to this abiotic factor due to its tropical evolutionary history and adaptations to temperate climates. Temperature is known to affect odonate physiology including life-history traits such as developmental rate, phenology and seasonal regulation as well as immune function and the production of pigment for thermoregulation. A range of behaviours are likely to be affected which will, in turn, influence other parts of the aquatic ecosystem, primarily through trophic interactions. Temperature may influence changes in geographical distributions, through a shifting of species' fundamental niches, changes in the distribution of suitable habitat and variation in the dispersal ability of species. Finally, such a rapid change in the environment results in a strong selective pressure towards adaptation to cope and the inevitable loss of some populations and, potentially, species. Where data are lacking for odonates, studies on other invertebrate groups will be considered. Finally, directions for research are suggested, particularly laboratory studies that investigate underlying causes of climate-driven macroecological patterns

Evolution of sex-specific pace-of-life syndromes: genetic architecture and physiological mechanisms

Sex differences in life history, physiology, and behavior are nearly ubiquitous across taxa, owing to sex-specific selection that arises from different reproductive strategies of the sexes. The pace-of-life syndrome (POLS) hypothesis predicts that most variation in such traits among individuals, populations, and species falls along a slow-fast pace-of-life continuum. As a result of their different reproductive roles and environment, the sexes also commonly differ in pace-of-life, with important consequences for the evolution of POLS. Here, we outline mechanisms for how males and females can evolve differences in POLS traits and in how such traits can covary differently despite constraints resulting from a shared genome. We review the current knowledge of the genetic basis of POLS traits and suggest candidate genes and pathways for future studies. Pleiotropic effects may govern many of the genetic correlations, but little is still known about the mechanisms involved in trade-offs between current and future reproduction and their integration with behavioral variation. We highlight the importance of metabolic and hormonal pathways in mediating sex differences in POLS traits; however, there is still a shortage of studies that test for sex specificity in molecular effects and their evolutionary causes. Considering whether and how sexual dimorphism evolves in POLS traits provides a more holistic framework to understand how behavioral variation is integrated with life histories and physiology, and we call for studies that focus on examining the sex-specific genetic architecture of this integration

Functional correlates of skull shape in Chiroptera: feeding and echolocation adaptations.

Morphological, functional and behavioural adaptations of bats are among the most diverse within mammals. A strong association between bat skull morphology and feeding behaviour has been suggested previously. However, morphological variation related to other drivers of adaptation, in particular echolocation, remains understudied. We assessed variation in skull morphology with respect to ecology (diet and emission type) and function (bite force, masticatory muscles and echolocation characteristics) using geometric morphometrics and comparative methods. Our study suggests that variation in skull shape of 10 bat families is the result of adaptations to broad dietary categories and sound emission types (oral or nasal). Skull shape correlates with echolocation parameters only in a subsample of insectivorous species, possibly because they (almost) entirely rely on this sensory system for locating and capturing prey. Insectivores emitting low frequencies are characterised by a ventrally tilted rostrum, a trait not associated with feeding parameters. This result questions the validity of a trade-off between feeding and echolocation function. Our study advances understanding of the relationship between skull morphology and specific features of echolocation and suggests that evolutionary constraints due to echolocation may differ between different groups within the Chiroptera