Closely related species show species-specific environmental responses and different spatial conservation needs: Prionailurus cats in the Indian subcontinent.

Phylogenetically closely related species are often assumed to have similar responses to environmental conditions, but species-specific responses have also been described. These two scenarios may have different conservation implications. We tested these two hypotheses for Prionailurus cats (P. rubiginosus, P. bengalensis, P. viverrinus) in the Indian subcontinent and show its implications on species current protected area coverage and climatic suitability trends through time. We fitted ecological niche models with current environmental conditions and calculated niche overlap. In addition, we developed a model for the Jungle Cat Felis chaus to compare species responses and niche overlap estimates within Prionailurus with those for a related sympatric small cat species. Then we estimated the proportion of current suitable environment covered by protected area and projected climatic models from past (last interglacial) to future (2070; RCP4.5 and RCP8.5) conditions to show implications on population management and conservation. The hypothesis of a similar response and niche overlap among closely related species is not supported. Protected area coverage was lowest for P. viverrinus (mean = 0.071, SD = 0.012) and highest for P. bengalensis (mean = 0.088, SD = 0.006). In addition, the proportion of the subcontinent with suitable climate varied through time and was species-specific. For P. bengalensis, climatic suitability shrunk since at least the mid-Holocene, a trend that can be intensified by human-induced climate warming. Concerning P. viverrinus, most predictions show stable future climatic suitability, but a few indicated potential loss. Climatic suitability for P. rubiginous was predicted to remain stable but the species exhibited a negative association with intensive agriculture. Similar responses to environmental change by phylogenetically closely related species should not be assumed and have implications on protected area coverage and natural trends of species climatic suitability over time. This should be taken into account during conservation and management actions.

Be careful with your principal components.

Principal components analysis (PCA) is a common method to summarize a larger set of correlated variables into a smaller and more easily interpretable axes of variation. However, the different components need to be distinct from each other to be interpretable otherwise they only represent random directions. This is a fundamental assumption of PCA and, thus, needs to be tested every time. Sample correlation matrices will always result in a pattern of decreasing eigenvalues even if there is no structure. Tests are, therefore, needed to discern real patterns from illusionary ones. Furthermore, the loadings of the vectors need to be larger than expected by random data to be useful in the calculation of PC-scores. PC-scores calculated from nondistinct PC's have very large standard errors and cannot be used for biological interpretations. I give a number of examples to illustrate the potential problems with PCA. Robustness of the PC's increases with increasing sample size but not with the number of traits. I review a few simple test statistics appropriate for testing PC's and use a real-world example to illustrate how this can be done using randomization tests. PCA can be very useful but great care is needed to avoid spurious results.

Lamarck, the Father of Evolutionary Ecology?

Lamarck realized life had evolved from simple to more complex forms, due to adaptation to a changing environment over time. Though he was wrong in many details, he got the overall picture right. Thus, he can be seen as the first evolutionary ecologist, connecting evolutionary change in organisms to their environment.

Rapid evolution of parasite resistance in a warmer environment: insights from a large scale field experiment.

Global climate change is expected to have major effects on host-parasite dynamics, with potentially enormous consequences for entire ecosystems. To develop an accurate prognostic framework, theoretical models must be supported by empirical research. We investigated potential changes in host-parasite dynamics between a fish parasite, the eyefluke Diplostomum baeri, and an intermediate host, the European perch Perca fluviatilis, in a large-scale semi-enclosed area in the Baltic Sea, the Biotest Lake, which since 1980 receives heated water from a nuclear power plant. Two sample screenings, in two consecutive years, showed that fish from the warmer Biotest Lake were now less parasitized than fish from the Baltic Sea. These results are contrasting previous screenings performed six years after the temperature change, which showed the inverse situation. An experimental infection, by which perch from both populations were exposed to D. baeri from the Baltic Sea, revealed that perch from the Baltic Sea were successfully infected, while Biotest fish were not. These findings suggest that the elevated temperature may have resulted, among other outcomes, in an extremely rapid evolutionary change through which fish from the experimental Biotest Lake have gained resistance to the parasite. Our results confirm the need to account for both rapid evolutionary adaptation and biotic interactions in predictive models, and highlight the importance of empirical research in order to validate future projections.

The stability of the G-matrix: The role of spatial heterogeneity.

The temporal stability of the genetic variance-covariance matrix (G) has been discussed for a long time in the evolutionary literature. A common assumption in all studies, including empirical ones, is that spatial heterogeneity is minor such that the population can be represented by a single mean and variance. We use the well-established allocation-acquisition model to analyze the effect of relaxing of this assumption, simulating a case where the population is divided into patches with a variance in quality between patches. This variance can in turn differ between years. We found that changes in spatial variance in patch quality over years can make the G-matrix vary substantially over years and that the estimated genetic correlations, evolvability, and response to selection are different dependent on whether spatial heterogeneity is taken into account or not. This will have profound implications for our ability to predict evolutionary change and understanding of the evolutionary process.

Isolation over 35 years in a heated biotest basin causes selection on MHC class IIß genes in the European perch (Perca fluviatilis L.).

Genes that play key roles in host immunity such as the major histocompatibility complex (MHC) in vertebrates are expected to be major targets of selection. It is well known that environmental conditions can have an effect on host-parasite interactions and may thus influence the selection on MHC. We analyzed MHC class IIß variability over 35 years in a population of perch (Perca fluviatilis) from the Baltic Sea that was split into two populations separated from each other. One population was subjected to heating from cooling water of a nuclear power plant and was isolated from the surrounding environment in an artificial lake, while the other population was not subjected to any change in water temperature (control). The isolated population experienced a change of the allelic composition and a decrease in allelic richness of MHC genes compared to the control population. The two most common MHC alleles showed cyclic patterns indicating ongoing parasite-host coevolution in both populations, but the alleles that showed a cyclic behavior differed between the two populations. No such patterns were observed at alleles from nine microsatellite loci, and no genetic differentiation was found between populations. We found no indications for a genetic bottleneck in the isolated population during the 35 years. Additionally, differences in parasitism of the current perch populations suggest that a change of the parasite communities has occurred over the isolation period, although the evidence in form of in-depth knowledge of the change of the parasite community over time is lacking. Our results are consistent with the hypothesis of a selective sweep imposed by a change in the parasite community.

Increase in body size is correlated to warmer winters in a passerine bird as inferred from time series data.

Climate change is expected to affect natural populations in many ways. One way of getting an understanding of the effects of a changing climate is to analyze time series of natural populations. Therefore, we analyzed time series of 25 and 20 years, respectively, in two populations of the citril finch (Carduelis citrinella) to understand the background of a dramatic increase in wing length in this species over this period, ranging between 1.3 and 2.9 phenotypic standard deviations. We found that the increase in wing length is closely correlated to warmer winters and in one case to rain in relation to temperature in the summer. In order to understand the process of change, we implemented seven simulation models, ranging from two nonadaptive models (drift and sampling), and five adaptive models with selection and/or phenotypic plasticity involved and tested these models against the time series of males and females from the two population separately. The nonadaptive models were rejected in each case, but the results were mixed when it comes to the adaptive models. The difference in fit of the models was sometimes not significant indicating that the models were not different enough. In conclusion, the dramatic change in mean wing length can best be explained as an adaptive response to a changing climate.

The importance of selection at the level of the pair over 25 years in a natural population of birds.

Knowledge of the pattern of selection in natural populations is fundamental for our understanding of the evolutionary process. Selection at higher levels has gained considerable theoretical support in recent years, and one possible level of selection is the breeding pair where fitness is a function of the pair and cannot be reduced to single individuals. We analyzed the importance of pair-level selection over 25 years in a natural population of the collared flycatcher. Pair-level selection was significant in five and probably in another 9 years. The relative importance of pair-level selection varied over years and can have stronger or the same strength as directional selection. This means that selection can act on the combination of the breeding pair in addition to selection on each individual separately. Overall, the conservative estimates obtained here show that this is a potentially important form of selection.

Integration of a hand-reared chimpanzee (Pan troglodytes) infant into a social group of conspecifics.

Rejections of infants among non-human primates occasionally occur in the wild as well as in captive settings. Controlled adoptions of orphans and introductions of individuals into new groups are therefore sometimes necessary in captivity. Consequently, behavioral research on integration procedures and on the acceptance of infants by adoptive mothers is much needed. In this study, the introduction and subsequent adoption were examined in an 18-month-old hand-reared chimpanzee (Pan troglodytes). The infant was introduced into an age/sex-diversified social group of conspecifics at Furuvik Zoo, Gävle, Sweden, and continuous focal data was collected during the final stage of integration, including infant care exhibited by the group members and the infant's secure base behavior. The infant was successfully integrated into the group and engaged in positive social interactions with all group members. An adult primiparous female chimpanzee formed a bond resembling a mother-infant relationship with the infant, which continues to be maintained at publication. However, the female initially showed very limited interest in the infant. It was, in fact, two other younger female group members that exhibited most infant care. The infant's secure base behavior patterns indicate that she adapted well to the new circumstances in the chimpanzee group as the integration progressed. This provides evidence that a final adopter does not necessarily initially show maternal interest and that there can be flexibility in maternal behavior in adult chimpanzee females. Moreover, the methods applied employing gradual familiarization with all the group members and the use of an integration enclosure, may have contributed to a successful result. These findings extend our knowledge of introduction procedures in captivity as well as provide information on foster mother-infant attachment in chimpanzees.

How often do they meet? Genetic similarity between European populations of a potential disease vector Culex pipiens.

BACKGROUND: Species in the Culex pipiens complex are common almost all over the world and represent important vectors for many serious zoonotic diseases. Even if, at the moment, many of the pathogens potentially transmitted by Cx. pipiens are not a problem in northern Europe, they may, with increasing temperatures and changing ecosystems caused by climate change, move northward in the future. Therefore, the question whether or not the Cx. pipiens populations in northern Europe will be competent vectors for them is of high importance. One way to estimate the similarity and the rate of contact between European Cx. pipiens populations is to look at the gene exchange between these populations. METHODS: To test the genetic diversity and degree of differentiation between European Cx. pipiens populations, we used eight microsatellite markers in 10 mosquito populations originating from northern, central, and southern Europe. RESULTS: We found that three of the analyzed populations were very different from the rest of the populations and they also greatly differed from each other. When these three populations were removed, the variance among the rest of the populations was low, suggesting an extensive historic gene flow between many European Cx. pipiens populations. CONCLUSIONS: This suggests that infectious diseases spread by this species may not be associated with a certain vector genotype but rather with suitable environmental conditions. Consequently, we would expect these pathogens to disperse northward with favorable climatic parameters.

Variation in predator species abundance can cause variable selection pressure on warning signaling prey.

Predation pressure is expected to drive visual warning signals to evolve toward conspicuousness. However, coloration of defended species varies tremendously and can at certain instances be considered as more camouflaged rather than conspicuous. Recent theoretical studies suggest that the variation in signal conspicuousness can be caused by variation (within or between species) in predators' willingness to attack defended prey or by the broadness of the predators' signal generalization. If some of the predator species are capable of coping with the secondary defenses of their prey, selection can favor reduced prey signal conspicuousness via reduced detectability or recognition. In this study, we combine data collected during three large-scale field experiments to assess whether variation in avian predator species (red kite, black kite, common buzzard, short-toed eagle, and booted eagle) affects the predation pressure on warningly and non-warningly colored artificial snakes. Predation pressure varied among locations and interestingly, if common buzzards were abundant, there were disadvantages to snakes possessing warning signaling. Our results indicate that predator community can have important consequences on the evolution of warning signals. Predators that ignore the warning signal and defense can be the key for the maintenance of variation in warning signal architecture and maintenance of inconspicuous signaling.

Leptin, a tool of parasites?

One common physiological phenomenon that is involved both in infectious and in malignant processes is the reduction in appetite: disease anorexia. An increase in plasma levels of leptin with inflammation is thought to be involved in this process. However, from an evolutionary perspective, in certain cases, it would be more adaptive for an internal parasite to stimulate the appetite of the host instead of causing its suppression. We tested whether a parasitic infection with the larvae of the helminth parasite Taenia taeniaformis affects the levels of appetite-regulating proteins, such as leptin, ghrelin and neuropeptide-Y (NPY) in wild yellow-necked mouse (Apodemus flavicollis). We found that infected mice had lower plasma levels of leptin and increased levels of NPY than the uninfected subjects. Ghrelin levels were not associated with the occurrence of the parasites; however, these levels strongly correlated with the levels of NPY. This study suggests a possible manipulation by parasitic larvae of appetite regulation in infected subjects.

Sex-specific genetic variances in life-history and morphological traits of the seed beetle Callosobruchus maculatus.

Knowledge of heritability and genetic correlations are of central importance in the study of adaptive trait evolution and genetic constraints. We use a paternal half-sib-full-sib breeding design to investigate the genetic architecture of three life-history and morphological traits in the seed beetle, Callosobruchus maculatus. Heritability was significant for all traits under observation and genetic correlations between traits (r(A)) were low. Interestingly, we found substantial sex-specific genetic effects and low genetic correlations between sexes (r(MF)) in traits that are only moderately (weight at emergence) to slightly (longevity) sexually dimorphic. Furthermore, we found an increased sire ([Formula: see text]) compared to dam ([Formula: see text]) variance component within trait and sex. Our results highlight that the genetic architecture even of the same trait should not be assumed to be the same for males and females. Furthermore, it raises the issue of the presence of unnoticed environmental effects that may inflate estimates of heritability. Overall, our study stresses the fact that estimates of quantitative genetic parameters are not only population, time, environment, but also sex specific. Thus, extrapolation between sexes and studies should be treated with caution.

If F(ST) does not measure neutral genetic differentiation, then comparing it with Q(ST) is misleading. Or is it?

The comparison between neutral genetic differentiation (F(ST) ) and quantitative genetic differentiation (Q(ST) ) is commonly used to test for signatures of selection in population divergence. However, there is an ongoing discussion about what F(ST) actually measures, even resulting in some alternative metrics to express neutral genetic differentiation. If there is a problem with F(ST) , this could have repercussions for its comparison with Q(ST) as well. We show that as the mutation rate of the neutral marker increases, F(ST) decreases: a higher within-population heterozygosity (He) yields a lower F(ST) value. However, the same is true for Q(ST) : a higher mutation rate for the underlying QTL also results in a lower Q(ST) estimate. The effect of mutation rate is equivalent in Q(ST) and F(ST) . Hence, the comparison between Q(ST) and F(ST) remains valid, if one uses neutral markers whose mutation rates are not too high compared to those of quantitative traits. Usage of highly variable neutral markers such as hypervariable microsatellites can lead to serious biases and the incorrect inference that divergent selection has acted on populations. Much of the discussion on F(ST) seems to stem from the misunderstanding that it measures the differentiation of populations, whereas it actually measures the fixation of alleles. In their capacity as measures of population differentiation, Hedrick's G'(ST) and Jost's D reach their maximum value of 1 when populations do not share alleles even when there remains variation within populations, which invalidates them for comparisons with Q(ST) .

Can preference for oviposition sites initiate reproductive isolation in Callosobruchus maculatus?

Theory has identified a variety of evolutionary processes that may lead to speciation. Our study includes selection experiments using different host plants and test key predictions concerning models of speciation based on host plant choice, such as the evolution of host use (preference and performance) and assortative mating. This study shows that after only ten generations of selection on different resources/hosts in allopatry, strains of the seed beetle Callosobruchus maculatus develop new resource preferences and show resource-dependent assortative mating when given the possibility to choose mates and resources during secondary contact. The resulting reduced gene flow between the different strains remained for two generations after contact before being overrun by disassortative mating. We show that reduced gene flow can evolve in a population due to a link between host preference and assortative mating, although this result was not found in all lines. However, consistent with models of speciation, assortative mating alone is not sufficient to maintain reproductive isolation when individuals disperse freely between hosts. We conclude that the evolution of reproductive isolation in this system cannot proceed without selection against hybrids. Other possible factors facilitating the evolution of isolation would be longer periods of allopatry, the build up of local adaptation or reduced migration upon secondary contact.

Effects of chronic leptin administration on nitric oxide production and immune responsiveness of greenfinches.

Leptin and nitric oxide (NO) are both important messengers in intra- and intercellular communication systems in vertebrates. Several studies have demonstrated an involvement of both substances in the immune response. Here we tested the effects of chronic leptin and anti-leptin treatments on the NO production and phytohaemagglutinin- (PHA) induced cutaneous inflammatory response in a wild passerine, the greenfinch (Carduelis chloris). Plasma leptin levels of individual birds were consistent in time but could be still temporarily increased by administration of recombinant chicken leptin. Increase of plasma leptin was also induced by administration of anti-leptin, which can be most likely explained by increased endogenous leptin production due to disruption of signalling pathways. Contrary to previous findings in mammals, leptin administration reduced systemic NO production. Leptin increased cutaneous swelling response to PHA. This immune-enhancing effect was observable despite the similar plasma leptin levels of leptin-treated and control birds at the time of measurement of immune responses, i.e., 9 days after start of the treatments. This provides evidence for a delayed or long-term potentiation of the cells and cytokines involved. The effects of leptin administration on NO production and immune responsiveness were age-dependent, which indicates the complexity of underlying regulatory mechanisms.

Genotype-temperature interaction in the regulation of development, growth, and morphometrics in wild-type, and growth-hormone transgenic coho salmon.

BACKGROUND: The neuroendocrine system is an important modulator of phenotype, directing cellular genetic responses to external cues such as temperature. Behavioural and physiological processes in poikilothermic organisms (e.g. most fishes), are particularly influenced by surrounding temperatures. METHODOLOGY/PRINCIPAL FINDINGS: By comparing the development and growth of two genotypes of coho salmon (wild-type and transgenic with greatly enhanced growth hormone production) at six different temperatures, ranging between 8 degrees and 18 degrees C, we observed a genotype-temperature interaction and possible trend in directed neuroendocrine selection. Differences in growth patterns of the two genotypes were compared by using mathematical models, and morphometric analyses of juvenile salmon were performed to detect differences in body shape. The maximum hatching and alevin survival rates of both genotypes occurred at 12 degrees C. At lower temperatures, eggs containing embryos with enhanced GH production hatched after a shorter incubation period than wild-type eggs, but this difference was not apparent at and above 16 degrees C. GH transgenesis led to lower body weights at the time when the yolk sack was completely absorbed compared to the wild genotype. The growth of juvenile GH-enhanced salmon was to a greater extent stimulated by higher temperatures than the growth of the wild-type. Increased GH production significantly influenced the shape of the salmon growth curves. CONCLUSIONS: Growth hormone overexpression by transgenesis is able to stimulate the growth of coho salmon over a wide range of temperatures. Temperature was found to affect growth rate, survival, and body morphology between GH transgenic and wild genotype coho salmon, and differential responses to temperature observed between the genotypes suggests they would experience different selective forces should they ever enter natural ecosystems. Thus, GH transgenic fish would be expected to differentially respond and adapt to shifts in environmental conditions compared with wild type, influencing their ability to survive and interact in ecosystems. Understanding these relationships would assist environmental risk assessments evaluating potential ecological effects.

Adaptive dynamics of regulatory networks: size matters.

To accomplish adaptability, all living organisms are constructed of regulatory networks on different levels which are capable to differentially respond to a variety of environmental inputs. Structure of regulatory networks determines their phenotypical plasticity, that is, the degree of detail and appropriateness of regulatory replies to environmental or developmental challenges. This regulatory network structure is encoded within the genotype. Our conceptual simulation study investigates how network structure constrains the evolution of networks and their adaptive abilities. The focus is on the structural parameter network size. We show that small regulatory networks adapt fast, but not as good as larger networks in the longer perspective. Selection leads to an optimal network size dependent on heterogeneity of the environment and time pressure of adaptation. Optimal mutation rates are higher for smaller networks. We put special emphasis on discussing our simulation results on the background of functional observations from experimental and evolutionary biology.

The species status of the Corsican finch Carduelis corsicana assessed by three genetic markers with different rates of evolution.

Citril finches Carduelis c. citrinella and Corsican finches Carduelis c. corsicana represent two closely related forms, endemic to European mountains and some Mediterranean Islands. Their taxonomic status has recently been disputed based on the results from mitochondrial genes. We show that the use of different genetic markers may lead to different results in these two putative species. Using the mitochondrial ATP8+6 we found a clear difference between C. c. citrinella and C. c. corsicana (3.2%) suggesting a divergence time of 1.2MYR. In contrast, no clear difference was found using two nuclear genes. The mismatch between the different markers suggests that the separation of C. c. citrinella and C. c. corsicana is likely to be a rather recent event, involving bottlenecks, which have enhanced the divergence in the mtDNA. Our results call for caution when using mtDNA alone for assessing times of divergence and rates of evolution. We did not find any support for the view that C. c. corsicana is more related to Carduelis carduelis than C. c. citrinella as proposed by previous studies.

Quantitative trait evolution and environmental change.

BACKGROUND: Given the recent changes in climate, there is an urgent need to understand the evolutionary ability of populations to respond to these changes. METHODOLOGY/PRINCIPAL FINDINGS: We performed individual-based simulations with different shapes of the fitness curve, different heritabilities, different levels of density compensation, and different autocorrelation of environmental noise imposed on an environmental trend to study the ability of a population to adapt to changing conditions. The main finding is that when there is a positive autocorrelation of environmental noise, the outcome of the evolutionary process is much more unpredictable compared to when the noise has no autocorrelation. In addition, we found that strong selection resulted in a higher load, and more extinctions, and that this was most pronounced when heritability was low. The level of density-compensation was important in determining the variance in load when there was strong selection, and when genetic variance was lower when the level of density-compensation was low. CONCLUSIONS: The strong effect of the details of the environmental fluctuations makes predictions concerning the evolutionary future of populations very hard to make. In addition, to be able to make good predictions we need information on heritability, fitness functions and levels of density compensation. The results strongly suggest that patterns of environmental noise must be incorporated in future models of environmental change, such as global warming.