Ecological divergence in the presence of gene flow in two closely related Oryza species (Oryza rufipogon and O. nivara)

Ecological divergence plays a prominent role in the process of speciation, but how divergence occurs in the face of gene flow is still less clear, and remains controversial among evolutionists. Here we investigated the nucleotide diversity, divergence and gene flow between Oryza nivara and O. rufipogon using sequences of seven chloroplast and nuclear loci. By analysing samples from 26 wild populations across the geographic ranges of the two species, we showed that both species were highly structured and O. rufipogon maintained a higher level of species‐wide diversity than O. nivara. Notably, phylogenetic, amova and F_ST analyses were unable to detect significant nucleotide differentiation between the two species. We estimated that the two species began to diverge at c. 0.16 million years ago. Our coalescent‐based simulations strongly rejected the simple isolation model of zero migration between species, but rather provided unambiguous evidence of bidirectional gene flow between species, particularly from O. rufipogon to O. nivara. Our simulations also indicated that gene flow was recurrent during the divergence process rather than arising from secondary contact after allopatric divergence. In conjunction with different morphological and life‐history traits and habitat preference in the two species, this study supports the hypothesis that these Oryza species are better treated as ecotypes that diverged quite recently and are still under the process of divergence. Importantly, we demonstrate the ecological divergence between O. rufipogon and O. nivara in the presence of significant gene flow, implying that natural selection plays a primary role in driving the divergence of the two Oryza species.     

Life‐history strategies indicate live‐bearing in Nothosaurus (Sauropterygia)

In Sauropterygia, a diverse group of Mesozoic marine reptiles, fossil evidence of viviparity (live‐bearing) only exists for Pachypleurosauria and Plesiosauria, and was assumed to also be the case for nothosaurs. Previous studies have successfully applied an extant squamate model to sauropterygian life‐history traits. In extant squamates, oviparity and viviparity are associated with differences in life‐history trait combinations. We establish growth curves for Nothosaurus specimens based on their humeral histology. We then analyse life‐history traits derived from these curves and compare inferred traits to those of modern squamates and pachypleurosaurs to assess their reproduction mode. We show that birth to adult size ratios (i.e. birth size divided by the mother's size) provide good estimates of clutch sizes in extant squamates and in viviparous extinct marine reptiles, but these ratios cannot discriminate viviparous and oviparous squamates. Thus, large ratios do not indicate viviparity in fossil taxa to which the extant squamate model is applicable. Applying differences in birth size, age at maturation, and maximum longevity that are observed between extant viviparous and oviparous squamates to our Nothosaurus sample, we identified 7 out of 24 specimens as being potentially viviparous. Conversely, they suggested oviparity for many nothosaurs but also for many pachypleurosaur samples. Under the assumption that the entire clade Pachypleurosauria was viviparous, the majority of nothosaurs would also have been viviparous as they comprised trait combinations similar to those seen in pachypleurosaurs. Overall, this suggests that within nothosaurs and pachypleurosaurs both reproduction modes existed in different taxa.     

Evolution of biometric and life‐history traits in lizards (Gallotia) from the Canary Islands

The aim was to study as to how biometric and life‐history traits of endemic lacertids in the Canary Islands (genus Gallotia) may have evolved, and possible factors affecting the diversification process of this taxon on successively appearing islands have been deduced. To that end, comparative analyses of sexual dimorphism and scaling of different body, head and life‐history traits to body size in 10 species/subspecies of Gallotia have been carried out. Both Felsenstein's independent contrasts and Huey and Bennett's ‘minimum evolution’ analyses show that male and female snout‐vent length (SVL) changed proportionally (sexual size dimorphism not changing with body size) throughout the evolution of these lizards and all within‐sex biometric traits have changed proportionally to SVL. Life‐history traits (size at sexual maturity, clutch size, hatchling SVL and mass, and life span) are highly correlated with adult female body size, the first two being the only traits with a positive allometry to female SVL. These results, together with the finding that the slope of hatchling SVL to female SVL regression was lower than that of SVL at maturity to female SVL, indicates that larger females reach maturity at a larger size, have larger clutches and, at the same time, have relatively smaller hatchlings than smaller females. There was no significant correlation between any pair of life‐history traits after statistically removing the effect of body size. As most traits changed proportionally to SVL, the major evolutionary change has been that of body size (a ca. threefold change between the largest and the smallest species), that is suggested to be the effect of variable ecological conditions faced by founder lizards in each island.     

Life‐history traits and physiological limits of the alpine fly Drosophila nigrosparsa (Diptera: Drosophilidae): A comparative study

Interspecific variation in life‐history traits and physiological limits can be linked to the environmental conditions species experience, including climatic conditions. As alpine environments are particularly vulnerable under climate change, we focus on the montane‐alpine fly Drosophila nigrosparsa. Here, we characterized some of its life‐history traits and physiological limits and compared these with those of other drosophilids, namely Drosophila hydei, Drosophila melanogaster, and Drosophila obscura. We assayed oviposition rate, longevity, productivity, development time, larval competitiveness, starvation resistance, and heat and cold tolerance. Compared with the other species assayed, D. nigrosparsa is less fecund, relatively long‐living, starvation susceptible, cold adapted, and surprisingly well heat adapted. These life‐history characteristics provide insights into invertebrate adaptations to alpine conditions which may evolve under ongoing climate change.     

The effects of selection,drift and genetic variation on life‐history trait divergence among insular populations of natterjack toad,Bufo calamita

Although loss of genetic variation is frequently assumed to be associated with loss of adaptive potential, only few studies have examined adaptation in populations with little genetic variation. On the Swedish west coast, the northern fringe populations of the natterjack toad Bufo calamita inhabit an atypical habitat consisting of offshore rock islands. There are strong among‐population differences in the amount of neutral genetic variation, making this system suitable for studies on mechanisms of trait divergence along a gradient of within‐population genetic variation. In this study, we examined the mechanisms of population divergence using Q_ST–F_ST comparisons and correlations between quantitative and neutral genetic variation. Our results suggest drift or weak stabilizing selection across the six populations included in this study, as indicated by low Q_ST–F_ST values, lack of significant population × temperature interactions and lack of significant differences among the islands in breeding pond size. The six populations included in this study differed in both neutral and quantitative genetic variation. Also, the correlations between neutral and quantitative genetic variation tended to be positive, however, the relatively small number of populations prevents any strong conclusions based on these correlations. Contrary to the majority of Q_ST–F_ST comparisons, our results suggest drift or weak stabilizing selection across the examined populations. Furthermore, the low heritability of fitness‐related traits may limit evolutionary responses in some of the populations.     

Genetic variation in the life‐history traits of Epiphyas postvittana: population structure and local adaptation

The light brown apple moth, Epiphyas postvittana (Walker) shows high intraspecific variability in morphological, physiological, demographic and behavioural characters. To gain insight into the extent of adaptation and evolutionary changes in response to environmental heterogeneity in this species, quantitative genetic analyses of life‐history variation were conducted for two natural populations under two thermal conditions (23°C and 28°C). Paternal half‐sib heritability and genetic correlation in six life‐history traits (i.e. development time, adult body weight, adult lifespan, age at first reproduction, the number of eggs laid during the first 5 days after emergence, and total fecundity) were compared. Significant heritabilities were shown consistently in development time; this is also true for adult body weight, except for the Canberra population at 23°C. However, neither population differences nor the effect of temperature were statistically detectable for any of these heritabilities, confirming the genetically determined flexibility. Positive genetic correlations between development time and adult body weight, and negative genetic correlations between the number of eggs laid during the first 5 days and adult lifespan were present for these populations at both temperatures, indicating the presence of genetic constraints. Pairwise comparisons of genetic correlations revealed the heterogeneity of the two populations and across temperatures. These results suggest that the structure of genetic covariance might have changed significantly during the divergence of natural populations and in response to the alteration of environmental conditions in E. postvittana.     

The predictability and magnitude of life‐history divergence to ecological agents of selection: a meta‐analysis in livebearing fishes

Environments causing variation in age‐specific mortality – ecological agents of selection – mediate the evolution of reproductive life‐history traits. However, the relative magnitude of life‐history divergence across selective agents, whether divergence in response to specific selective agents is consistent across taxa and whether it occurs as predicted by theory, remains largely unexplored. We evaluated divergence in offspring size, offspring number, and the trade‐off between these traits using a meta‐analysis in livebearing fishes (Poeciliidae). Life‐history divergence was consistent and predictable to some (predation, hydrogen sulphide) but not all (density, food limitation, salinity) selective agents. In contrast, magnitudes of divergence among selective agents were similar. Finally, there was a negative, asymmetric relationship between offspring‐number and offspring‐size divergence, suggesting greater costs of increasing offspring size than number. Ultimately, these results provide strong evidence for predictable and consistent patterns of reproductive life‐history divergence and highlight the importance of comparing phenotypic divergence across species and ecological selective agents.     

Life‐history trade‐offs under different larval diets in Drosophila suzukii (Diptera: Drosophilidae)

Most larval drosophilids eat the microorganisms that develop in rotting fruit, a relatively protein‐rich resource. By contrast, the spotted‐wing Drosophila suzukii Matsumara (Diptera: Drosophilidae) uniquely develops in ripening fruit, a protein‐poor, carbohydrate‐rich resource. This shift in larval nutritional niche has led to D. suzukii being a significant agricultural pest in the U.S.A. and Europe. Although occupying a new niche may benefit a species by reducing competition, adaptation in host use may generate trade‐offs affecting fitness. To test the hypothesis that fitness trade‐offs will change with adaptation to novel larval diets, D. suzukii larval development on either a diet of a fresh, ripe blueberry (a natural host) or standard artificial Drosophila media (protein‐rich) is compared and the effect of diet on development time from egg to adult, adult body size and male wing spot area, and female fecundity is assessed. Larval development time differs, with larvae on the blueberry emerging as adults earlier than those on the artificial medium, although other fitness measures do not vary between the two diets. In addition, the faster development time on a blueberry does not trade off with body size as expected, although early fecundity is delayed in females that develop on blueberries. Thus, adaptation to a novel larval diet environment does not come at a cost to the ability to develop in protein‐rich resources.     

A comparison of life history traits of sexual and asexual strains of the parasitoid wasp,Lysiphlebus fabarum (Braconidae: Aphidiinae)

1. This study examined biological characteristics of sexual and asexual strains of the parasitoid wasp, Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). 2. Strains were reared in different instar hosts (the black bean aphid, Aphis fabae Scopoli) under identical environmental conditions (21 °C, 65–75% RH, and LD 16:8 h). 3. Results showed that the second instar of the aphid is the most suitable growth stage for both strains, as the wasps that emerged from the second instar hosts were larger, more fecund, and had larger egg size. Trade‐offs between the fitness components of the parasitoid were clearer when the parasitoids were reared in suboptimal instars. 4. According to the results, sexual females emerged around 1 day earlier and lived around 0.5 day less than asexual females. Also, sexual females emerged with a lower initial egg load, although these wasps tend to have larger eggs than asexual females. Asexual females may enjoy greater longevity and higher developmental plasticity which suggests a higher degree of synchronization with pest population dynamism. 5. The results suggest that sexual wasps, in contrast to asexual wasps, invest more in egg size than in egg load. This study suggests strain‐specific adaptations of L. fabarum to different instars of the black bean aphid by which the allocation of nutritional resources to various functions differs between strains. 6. Furthermore, differences in life history traits between strains can greatly influence the population dynamics of each strain, and hence their effectiveness in suppressing pest populations.     

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.     

Spatial scale and divergent patterns of variation in adapted traits in the ocean

The geography of adaptive genetic variation is crucial to species conservation yet poorly understood in marine systems. We analyse the spatial scale of genetic variation in traits that broadly display adaptation throughout the range of a highly dispersive marine species. We conducted common garden experiments on the Atlantic silverside, Menidia menidia, from 39 locations along its 3000 km range thereby mapping genetic variation for growth rate, vertebral number and sex determination. Each trait displayed unique clinal patterns, with significant differences (adaptive or not) occurring over very small distances. Breakpoints in the cline differed among traits, corresponding only partially with presumed eco-geographical boundaries. Because clinal patterns are unique to each selected character, neutral genes or those coding for a single character cannot serve as proxies for the genetic structure as a whole. Conservation plans designed to protect essential genetic subunits of a species will need to account for such complex spatial structures.     

Divergent trends in life‐history traits between Atlantic salmon Salmo salar of wild and hatchery origin in the Baltic Sea

Four Atlantic salmon Salmo salar stocks in the Baltic Sea, varying in their breeding history, were studied for changes in life‐history traits over the years 1972–1995. Total length (L_T) at age of captured (L_TC) fish had increased throughout the study period, partly due to increased temperature and increased L_T at release, (L_TR) but also due to remaining cohort effects that could represent unaccounted environmental or genetic change. Simultaneously, maturation probabilities controlled for water temperature, L_TC and L_TR had increased in all stocks. The least change was observed in the River Tornionjoki S. salar that was subject only to supportive stockings originating from wild parents. These results suggest a long‐term divergence between semi‐natural and broodstock‐based S. salar stocks. Increased L_T at age explained advanced maturation only marginally, and it remains an open question to what extent the generally increased probabilities to mature at early age reflected underlying genetic changes.     

Natural and human‐mediated selection in a landrace of Thai rice (Oryza sativa)

Landrace rice in Thailand consists of managed populations grown under traditional and long‐standing agricultural practices. These populations evolve both in response to environmental conditions within the local agro‐ecosystem and in response to human activities. Single landraces are grown across varying environments and recently have experienced temporal changes in local environments due to climate change. Here we assess the interplay between natural selection in a changing climate and human‐mediated selection on the population genetic structure of Muey Nawng, a local landrace of Thai rice. Genetic diversity and population structure of landrace rice were assessed by a STRUCTURE analysis of 20 microsatellite loci. The first exon–intron junction of the waxy gene was sequenced to determine genotypes for glutinous or non‐glutinous grain starch. Muey Nawng rice is genetically variable and is structured based on starch grain types and the level of resistance to gall midge pest. A strong positive correlation was found between genetic diversity and the percentage of gall midge infestation. Variation in the waxy locus is correlated with starch quality; selection for non‐glutinous rice appears to involve additional genes. The dynamics of genetic diversity within Muey Nawng rice depends on three factors: (a) a genetic bottleneck caused by strong selection associated with gall midge infestation, (b) selection by local farmers for starch quality and (c) variation introduced by farmer practices for cultivation and seed exchange. These results, when taken in total, document the ability of landrace rice to quickly evolve in response to both natural and human‐mediated selection.     

Lethal and sub‐lethal effects of spinosad on the life‐history traits of army worm,Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae), and its fitness cost of resistance

Spodoptera litura is one of the most destructive polyphagous insect pests, with more than 120 host‐plant species. In our present study, a field‐collected population of S. litura when selected with spinosad for 11 consecutive generations resulted in the development of 3921‐fold resistance to spinosad as compared to the susceptible strain. The spinosad‐resistant strain of S. litura had a relatively high fitness cost (0.17) as compared to the susceptible strain. Furthermore, the lethal and sub‐lethal effects of different concentrations of spinosad were checked on the susceptible strain at different levels; i.e., LC₄₀, LC₃₀, LC₂₀ and LC₁₀, which revealed that the impact of spinosad on the life‐history traits of S. litura increased with the increase in concentration of spinosad. A significant impact of spinosad was recorded on the larval duration, pre‐pupal weight, pupal duration, pupal weight, reproductive potential and adult emergence. The outcomes of the current research clearly indicate that fitness cost of spinosad and its sub‐lethal effects have a significant impact on population dynamics of S. litura, for which it can be incorporated in integrated pest management.     

Within‐generation consequences of postsettlement mortality for trait composition in wild populations: An experimental test

There is a critical need to understand patterns and causes of intraspecific variation in physiological performance in order to predict the distribution and dynamics of wild populations under natural and human‐induced environmental change. However, the usual explanation for trait differences, local adaptation, fails to account for the small‐scale phenotypic and genetic divergence observed in fishes and other species with dispersive early life stages. We tested the hypothesis that local‐scale variation in the strength of selective mortality in early life mediates the trait composition in later life stages. Through in situ experiments, we manipulated exposure to predators in the coral reef damselfish Dascyllus aruanus and examined consequences for subsequent growth performance under common garden conditions. Groups of 20 recently settled D. aruanus were outplanted to experimental coral colonies in Moorea lagoon and either exposed to natural predation mortality (52% mortality in three days) or protected from predators with cages for three days. After postsettlement mortality, predator‐exposed groups were shorter than predator‐protected ones, while groups with lower survival were in better condition, suggesting that predators removed the longer, thinner individuals. Growth of both treatment groups was subsequently compared under common conditions. We did not detect consequences of predator exposure for subsequent growth performance: Growth over the following 37 days was not affected by the prior predator treatment or survival. Genotyping at 10 microsatellite loci did indicate, however, that predator exposure significantly influenced the genetic composition of groups. We conclude that postsettlement mortality did not have carryover effects on the subsequent growth performance of cohorts in this instance, despite evidence for directional selection during the initial mortality phase.     

Heritability of and strong single gene (Pgi) effects on life‐history traits in the Glanville fritillary butterfly

We estimated broad‐sense heritabilities (H²) of 13 female and seven male life‐history traits of the Glanville fritillary butterfly (Melitaea cinxia) under semi‐natural conditions in a large outdoor population cage. The analysis was based on full‐sib families collected as young larvae in the field and reared under common garden conditions. We found significant genetic variance in female lifespan, fecundity, number of matings and host‐plant preference as well as in male body mass and mobility. Apart from host‐plant preference, female traits that were more strongly correlated with lifetime reproductive success (LRS; measured as total number of eggs laid) had higher H². LRS itself exhibited significant heritability. Host‐plant preference had very high H², consistent with a previously reported genetically determined geographical cline in host‐plant preference in the study area. Lifespan and egg hatching rate were significantly associated with a SNP in the coding region of the Pgi gene, for which there is previous evidence for balancing selection. Selection on Pgi, which furthermore shows spatial and temporal variation, may maintain genetic variance in fitness‐related life‐history traits. In contrast, we found no strong evidence for life‐history trade‐offs.     

Environmental determinants of population divergence in life‐history traits for an invasive species: climate,seasonality and natural enemies

Invasive species cope with novel environments through both phenotypic plasticity and evolutionary change. However, the environmental factors that cause evolutionary divergence in invasive species are poorly understood. We developed predictions for how different life‐history traits, and plasticity in those traits, may respond to environmental gradients in seasonal temperatures, season length and natural enemies. We then tested these predictions in four geographic populations of the invasive cabbage white butterfly (Pieris rapae) from North America. We examined the influence of two rearing temperatures (20 and 26.7 °C) on pupal mass, pupal development time, immune function and fecundity. As predicted, development time was shorter and immune function was greater in populations adapted to longer season length. Also, phenotypic plasticity in development time was greater in regions with shorter growing seasons. Populations differed significantly in mean and plasticity of body mass and fecundity, but these differences were not associated with seasonal temperatures or season length. Our study shows that some life‐history traits, such as development time and immune function, can evolve rapidly in response to latitudinal variation in season length and natural enemies, whereas others traits did not. Our results also indicate that phenotypic plasticity in development time can also diverge rapidly in response to environmental conditions for some traits.