Degrees of change: between and within population variation in thermal reaction norms of phenology in a viviparous lizard

As the earth warms, populations will be faced with novel environments to which they may not be adapted. In the short term, populations can be buffered against the negative effects, or maximize the beneficial effects, of such environmental change via phenotypic plasticity and, in the longer term, via adaptive evolution. However, the extent and direction of these population-level responses will be dependent on the degree to which responses vary among the individuals within them (i.e., within population variation in plasticity), which is, itself, likely to vary among populations. Despite this, we have estimates of among-individual variation in plastic responses across multiple populations for only a few systems. This lack of data limits our ability to predict the consequences of environmental change for population and species persistence accurately. Here, we utilized a 16-yr data set from climatically distinct populations of the viviparous skink Niveoscincus ocellatus tracking over 1,200 litters from more than 600 females from each population to examine inter- and intrapopulation variability in the response of parturition date to environmental temperature. We found that these populations share a common population-mean reaction norm but differ in the degree to which reaction norms vary among individuals. These results suggest that even where populations share a common mean-level response, we cannot assume that they will be affected similarly by altered environmental conditions. If we are to assess how changing climates will impact species and populations accurately, we require estimates of how plastic responses vary both among and within populations.publishedVersio

Sex reversal explains some, but not all, climate-mediated sex ratio variation within a viviparous reptile

Evolutionary transitions in sex-determining systems have occurred frequently yet understanding how they occur remains a major challenge. In reptiles, transitions from genetic to temperature-dependent sex determination can occur if the gene products that determine sex evolve thermal sensitivity, resulting in sex-reversed individuals. However, evidence of sex reversal is limited to oviparous reptiles. Here we used thermal experiments to test whether sex reversal is responsible for differences in sex determination in a viviparous reptile, Carinascincus ocellatus, a species with XY sex chromosomes and population-specific sex ratio response to temperature. We show that sex reversal is occurring and that its frequency is related to temperature. Sex reversal was unidirectional (phenotypic males with XX genotype) and observed in both high- and low-elevation populations. We propose that XX-biased genotypic sex ratios could produce either male- or female-biased phenotypic sex ratios as observed in low-elevation C. ocellatus under variable rates of XX sex reversal. We discuss reasons why sex reversal may not influence sex ratios at high elevation. Our results suggest that the mechanism responsible for evolutionary transitions from genotypic to temperature-dependent sex determination is more complex than can be explained by a single process such as sex reversal

A comparative study on wear and corrosion behaviour of tungsten carbide-nickel and tungsten carbide-cobalt high velocity oxy-fuel (HVOF) for carbon steel blade

Nowadays, the demand of high wear and corrosion resistance of the components in various industry is increasing from time to time. Therefore, high velocity oxy-fuel (HVOF) thermal spray was introduced to protect machine components from wear and corrosion, to restore worn components and to improve the durability of the components. HVOF is one of the process of depositing a material layer over a base metal or substrate with characteristics of high flame velocity and moderate temperature. The main purpose of this present study is to characterize the structure of the tungsten carbide 10 wt.% nickel (WC-10Ni) and tungsten carbide 12 wt.% w cobalt (WC-12Co) coating deposited by means of HVOF thermal spray onto a continuous digester (CD) blade that made up from carbon steel. The morphology and chemical composition of the coating were characterized by scanning electron microscope (SEM), electron dispersive spectrometer (EDS), and x-ray diffraction (XRD). The hardness test was carried out by using Vickers micro-hardness tester with load of 490.3 mN (0.05 HV). The wear and corrosion behavior and mechanism for both coatings was compared. Three body wear test was carried out in term of weight loss and electrochemical test was performed in acidic media (mixture of sulfuric acid, H2SO4 and ilmenite) to obtain the corrosion rate of the coating. From the result, it shows that WC-12Co coating has finer grain size that is around 2.3 μm. WC-12Co has higher wear resistance due to high volume friction, low mean free path, high hardness and lower porosity distribution compared to WC-10Ni. Besides, the formation of secondary phase, W2C also affected the hardness of both coating, where this phase is harder than WC phase. For corrosion test, WC-12Co shows good corrosion resistance with small differences of corrison rate with WC-10Ni, that is only 0.7016 mm/y. As a conclusion, WC-12Co HVOF coating shows high potential on replacement of CD blade

Loss of genetic diversity and increased embryonic mortality in non-native lizard populations

Many populations are small and isolated with limited genetic variation and high risk of mating with close relatives. Inbreeding depression is suspected to contribute to extinction of wild populations, but the historical and demographic factors that contribute to reduced population viability are often difficult to tease apart. Replicated introduction events in non-native species can offer insights into this problem because they allow us to study how genetic variation and inbreeding depression are affected by demographic events (e.g. bottlenecks), genetic admixture and the extent and duration of isolation. Using detailed knowledge about the introduction history of 21 nonnative populations of the wall lizard Podarcis muralis in England, we show greater loss of genetic diversity (estimated from microsatellite loci) in older populations and in populations from native regions of high diversity. Loss of genetic diversity was accompanied by higher embryonic mortality in non-native populations, suggesting that introduced populations are sufficiently inbred to jeopardize long-term viability. However, there was no statistical correlation between population-level genetic diversity and average embryonic mortality. Similarly, at the individual level, there was no correlation between female heterozygosity and clutch size, infertility or hatching success, or between embryo heterozygosity and mortality. We discuss these results in the context of human-mediated introductions and how the history of introductions can play a fundamental role in influencing individual and population fitness in non-native species

Adaptive responses to cool climate promotes persistence of a non-native lizard

Successful establishment and range expansion of non-native species often require rapid accommodation of novel environments. Here, we use common-garden experiments to demonstrate parallel adaptive evolutionary response to a cool climate in populations of wall lizards (Podarcis muralis) introduced from southern Europe into England. Low soil temperatures in the introduced range delay hatching, which generates directional selection for a shorter incubation period. Non-native lizards from two separate lineages have responded to this selection by retaining their embryos for longer before oviposition-hence reducing the time needed to complete embryogenesis in the nest-and by an increased developmental rate at low temperatures. This divergence mirrors local adaptation across latitudes and altitudes within widely distributed species and suggests that evolutionary responses to climate can be very rapid. When extrapolated to soil temperatures encountered in nests within the introduced range, embryo retention and faster developmental rate result in one to several weeks earlier emergence compared with the ancestral state. We show that this difference translates into substantial survival benefits for offspring. This should promote short- and long-term persistence of non-native populations, and ultimately enable expansion into areas that would be unattainable with incubation duration representative of the native range

The diversity of population responses to environmental change

This is the final version. Available from Wiley via the DOI in this record.Data available from the Dryad Digital Repository: https:// doi.org/10.5061/dryad.d5f54s7The current extinction and climate change crises pressure us to predict population dynamics with ever-greater accuracy. Although predictions rest on the well-advanced theory of age-structured populations, two key issues remain poorly explored. Specifically, how the age-dependency in demographic rates and the year-to-year interactions between survival and fecundity affect stochastic population growth rates. We use inference, simulations and mathematical derivations to explore how environmental perturbations determine population growth rates for populations with different age-specific demographic rates and when ages are reduced to stages. We find that stage- vs. age-based models can produce markedly divergent stochastic population growth rates. The differences are most pronounced when there are survival-fecundity-trade-offs, which reduce the variance in the population growth rate. Finally, the expected value and variance of the stochastic growth rates of populations with different age-specific demographic rates can diverge to the extent that, while some populations may thrive, others will inevitably go extinct.Max Planck Society, Marie Curie FellowshipERCGerman Research FoundationSwiss National Science FoundationNational Science FoundationNational Institute of AgingRamon y Cajal Research GrantWenner-Gren FoundationLeakey FoundationNational Geographic SocietyZoological Society of San DiegoUniversity of PennsylvaniaArgentinean National Council of Researc

Female reproductive investment in response to male phenotype in wall lizards and its implications for introgression

The likelihood that females will breed or how much they invest in reproduction can depend on the characters of their male partners. Such differential allocation may enhance or limit gene flow between hybridizing lineages, in particular when the lineages have diverged in sexually selected characters. Populations of the common wall lizard (Podarcis muralis) in some regions of Italy exhibit striking exaggeration of coloration, morphology and behaviour compared to populations in France. The two forms hybridize in north-western Italy. Gene flow is directional and in concordance with the prediction that the exaggerated sexual characters of lizards of Italian origin give them a mating advantage. To evaluate if differential allocation contributes to asymmetric introgression, we tested experimentally if female reproductive investment is affected by male origin and male secondary sexual characters. Despite the large genetic and phenotypic divergence between males of Italian and French origin, females did not invest more when paired with males of the same origin, nor when paired with males with highly expressed sexual characters. Combined, these results suggest that female responses to male phenotypes in wall lizards are unimportant for explaining the directional pattern of gene flow in regions of secondary contact

Quo vadis amphibia? Global warming and breeding phenology in frogs, toads and salamanders

As the earth is getting warmer, many animals and plants have shifted their timing of breeding towards earlier dates. However, there is substantial variation between populations in phenological shifts that typically goes unexplained. Identification of the different location and species characteristics that drive such variable responses to global warming is crucial if we are to make predictions for how projected climate change scenarios will play out on local and global scales. Here we conducted a phylogenetically controlled meta-analysis of breeding phenology across frogs, toads and salamanders to examine the extent of variation in amphibian breeding phenology in response to global climate change. We show that there is strong geographic variation in response to global climate change, with species at higher latitudes exhibiting a more pronounced shift to earlier breeding than those at lower latitudes. Our analyses suggest that this latitude effect is a result of both the increased temperature (but not precipitation) at higher latitudes as well as a greater responsiveness by northern populations of amphibians to this change in temperature. We suggest that these effects should reinforce any direct effect of increasing warming at higher latitudes on breeding phenology. In contrast, we found very little contribution from other location factors or species traits. There was no evidence for a phylogenetic signal on advancing breeding phenology or responsiveness to temperature, suggesting that the amphibians that have been studied to date respond similarly to global warming

Are there benefits to being born asynchronously: an experimental test in a social lizard

Hatching asynchrony, or processes analogous to hatching asynchrony (i.e., birthing asynchrony), has now been identified in a number of nonavian taxa. These systems are of importance because, unlike birds, they allow us to decouple hatching asynchrony hypotheses related to adaptive hatching patterns, which suggest that it confers a fitness benefit to either the parent or offspring, from those that focus on the onset of parental incubation. However, to date research on these systems has remained descriptive, solely documenting the extent of asynchronous hatching/birthing. Here we provide the first experimental test of the potential adaptive nature of hatching/birthing asynchrony outside avian systems. Using the birth hormone arginine vasotocin, we manipulated the birthing (a)synchrony of females in a population of the lizard Egernia whitii and examined the effect on offspring growth and survival. We show offspring from asynchronous treatments suffered increased mortality but benefited from increased mass at 6 weeks compared with offspring in synchronous treatments. Differences in mortality and size between treatments were driven by offspring mass at birth, and the development of a greater mass hierarchy within asynchronous compared with synchronous litters. This resulted in the smaller offspring suffering both an increased risk of mortality and decreased growth. Despite this, the mechanism by which these patterns are produced remains unclear, as we found no link between birth order and mortality, size, or any factors that affected either of these. Copyright 2008, Oxford University Press.

Degrees of change: between and within population variation in thermal reaction norms of phenology in a viviparous lizard

As the earth warms, populations will be faced with novel environments to which they may not be adapted. In the short term, populations can be buffered against the negative effects, or maximize the beneficial effects, of such environmental change via phenotypic plasticity and, in the longer term, via adaptive evolution. However, the extent and direction of these population-level responses will be dependent on the degree to which responses vary among the individuals within them (i.e., within population variation in plasticity), which is, itself, likely to vary among populations. Despite this, we have estimates of among-individual variation in plastic responses across multiple populations for only a few systems. This lack of data limits our ability to predict the consequences of environmental change for population and species persistence accurately. Here, we utilized a 16-yr data set from climatically distinct populations of the viviparous skink Niveoscincus ocellatus tracking over 1,200 litters from more than 600 females from each population to examine inter- and intrapopulation variability in the response of parturition date to environmental temperature. We found that these populations share a common population-mean reaction norm but differ in the degree to which reaction norms vary among individuals. These results suggest that even where populations share a common mean-level response, we cannot assume that they will be affected similarly by altered environmental conditions. If we are to assess how changing climates will impact species and populations accurately, we require estimates of how plastic responses vary both among and within populations