Coadaptation of offspring begging and parental provisioning: A role for prenatal maternal effects?

Recent studies on birds have shown that offspring begging and parental provisioning covary at the phenotypic level, which is thought to reflect genetic correlations. However, prenatal maternal factors, like yolk testosterone, may also facilitate parent-offspring coadaptation via their effects on offspring begging and development. In fact, maternal effects are thought to adjust offspring phenotype to the environmental conditions they will experience after birth, which are in turn strongly dependent on the levels of parental provisioning. Using cross-fostering experiments in canaries, we tested the role of maternal effects on parent-offspring coadaptation from two different approaches. First, we analyzed whether females deposit yolk testosterone in relation to their own or their partner's prospective parental provisioning, measured as the rate of parental feeding to foster nestlings. Second, we investigated whether females deposit yolk testosterone in relation to costs they incurred when raising a previous brood, as this likely impinges on their capacity to provide parental care in the near future. However, from the results of both experiments we have no evidence that canary females deposit yolk testosterone in order to match offspring begging to the levels of care they and/or their partners provide. We therefore found no evidence that yolk testosterone facilitates parent-offspring coadaptation. In addition, our results suggest that the functional consequences of yolk testosterone deposition may relate to hatching asynchrony since it primarily varied with egg laying order.     

Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

Are offspring begging levels exaggerated beyond the parental optimum? Evidence from a bidirectional selection experiment

Parental care involves elaborate behavioural interactions between parents and their offspring, with offspring stimulating their parents via begging to provision resources. Thus, begging has direct fitness benefits as it enhances offspring growth and survival. It is nevertheless subject to a complex evolutionary trajectory, because begging may serve as a means for the offspring to manipulate parents in the context of evolutionary conflicts of interest. Furthermore, it has been hypothesized that begging is coadapted and potentially genetically correlated with parental care traits as a result of social selection. Further experiments on the causal processes that shape the evolution of begging are therefore essential. We applied bidirectional artificial selection on begging behaviour, using canaries (Serinus canaria) as a model species. We measured the response to selection, the consequences for offspring development, changes in parental care traits, here the rate of parental provisioning, as well as the effects on reproductive success. After three generations of selection, offspring differed in begging behaviour according to our artificial selection regime: nestlings of the high begging line begged significantly more than nestlings of the low begging line. Intriguingly, begging less benefitted the nestlings, as reflected by on average significantly higher growth rates, and increased reproductive success in terms of a higher number of fledglings in the low selected line. Begging could thus represent an exaggerated trait, possibly because parent–offspring conflict enhanced the selection on begging. We did not find evidence that we co‐selected on parental provisioning, which may be due to the lack of power, but may also suggest that the evolution of begging is probably not constrained by a genetic correlation between parental provisioning and offspring begging.     

Do parasites and antioxidant availability affect begging behaviour,growth rate and resistance to oxidative stress?

Early‐life trade‐offs faced by developing offspring can have long‐term consequences for their future fitness. Young offspring use begging displays to solicit resources from their parents and have been selected to grow fast to maximize survival. However, growth and begging behaviour are generally traded off against self‐maintenance. Oxidative stress, a physiological mediator of life‐history trade‐offs, may play a major role in this trade‐off by constraining, or being costly to, growth and begging behaviour. Yet, despite implications for the evolution of life‐history strategies and parent–offspring conflicts, the interplay between growth, begging behaviour and resistance to oxidative stress remains to be investigated. We experimentally challenged wild great tit (Parus major) offspring by infesting nests with a common ectoparasite, the hen flea (Ceratophyllus gallinae), and simultaneously tested for compensating effects of increased vitamin E availability, a common dietary antioxidant. We further quantified the experimental treatment effects on offspring growth, begging intensity and oxidative stress. Flea‐infested nestlings of both sexes showed reduced body mass during the first half of the nestling phase, but this effect vanished short before fledging. Begging intensity and oxidative stress of both sexes were unaffected by both experimental treatments. Feeding rates were not affected by the experimental treatments, but parents of flea‐infested nests fed nestlings with a higher proportion of caterpillars, the main source of antioxidants. Additionally, female nestlings begged significantly less than males in control nests, whereas both sexes begged at similar rates in vitamin E‐supplemented nests. Our study shows that a parasite exposure does not necessarily affect oxidative stress levels or begging intensity, but suggests that parents can compensate for negative effects of parasitism by modifying food composition. Furthermore, our results indicate that the begging capacity of the less competitive sex is constrained by antioxidant availability.     

Does landscape structure affect resource tracking by avian frugivores in a fragmented Afrotropical forest?

Frugivorous species heavily depend on patchy food resources and are believed to track these in space and time, thereby providing an important seed dispersal function that might be critical toward the regeneration of fruiting plants. However, isolation of suitable food patches due to habitat fragmentation or changes in landscape connectivity may hamper food tracking behaviour and adversely affect populations of both frugivores (through starvation) and food plants (through interruption of seed dispersal). We here test whether density fluctuations in four frugivorous Afrotropical bird species were larger and/or matched fluctuations in ripe fruit densities better in study plots embedded in large tracts of indigenous forest than in equally-sized plots embedded in cultivated lands. We compared these results with those of four non-frugivorous species (out-group) which were not expected to track fruit resources. Whereas densities of both frugivores and fruit crops strongly fluctuated in space and time, these fluctuations were not synchronised, nor did the level of synchrony differ in relation to matrix type. For some but not all bird species, lower densities and smaller temporal fluctuations in forest plots surrounded by cultivation may reflect decreased mobility. The observed fluctuations in bird densities most likely reflect exchange with the surrounding landscape matrix, suggesting that small pockets of fruiting trees in farmland may comprise critical food resources for frugivores inhabiting highly fragmented landscapes, apart from increasing connectivity for both bird and seed dispersal.     

Does a trade‐off between current reproductive success and survival affect the honesty of male signalling in species with male parental care?

Recent theory predicted that male advertisement will reliably signal investment in paternal care in species where offspring survival requires paternal care and males allocate resources between advertisement and care. However, the predicted relationship between care and advertisement depended on the marginal gains from investment in current reproductive traits. Life history theory suggests that these fitness gains are also subject to a trade‐off between current and future reproduction. Here, we investigate whether male signalling remains a reliable indicator of parental care when males allocate resources between current advertisement, paternal care and survival to future reproduction. We find that advertisement is predicted to remain a reliable signal of male care but that advertisement may cease to reliably indicate male quality because low‐quality males are predicted to invest in current reproduction, whereas higher‐quality males are able to invest in both current reproduction and survival to future reproduction.     

Does resource availability help determine the evolutionary route to multicellularity?

Genetic heterogeneity and homogeneity are associated with distinct sets of adaptive advantages and bottlenecks, both in developmental biology and population genetics. Whereas populations of individuals are usually genetically heterogeneous, most multicellular metazoans are genetically homogeneous. Observing that resource scarcity fuels genetic heterogeneity in populations, we propose that monoclonal development is compatible with the resource‐rich and stable internal environments that complex multicellular bodies offer. In turn, polyclonal development persists in tumors and in certain metazoans, both exhibiting a closer dependence on external resources. This eco‐evo‐devo approach also suggests that multicellularity may originally have emerged through polyclonal development in early metazoans, because of their reduced shielding from environmental fluctuations.     

Does fire affect the temporal pattern of trophic resource supply to pollinators and seed-dispersing frugivores in a Brazilian savanna community?

In tropical savannas, such as the campo cerrado in Brazil, fire plays an important role, affecting plant species’ life history. Since fire has the potential to modify the structure of savanna communities as a whole, it is expected that it may influence the resource supply for mutualists by altering the pattern of investment in sexual reproduction. We used an experimental approach to test if fire alters trophic resource availability to pollinators (nectar, pollen, and oil) and seed-dispersing frugivores (fleshy fruits) by altering the seasonality of reproductive phenophases in a savanna community. We sampled all individuals of 60 species that were common to both control and experimental fire treatments. Each month we recorded the number of reproductive individuals to test whether fire affected the temporal resource offered by the plant assemblage as a whole, and by each specific plant group supporting distinct groups of pollinators and seed-dispersing frugivores. We noticed that fire advanced the nectar, pollen, and fleshy fruit offered by the whole assemblage. Additionally, fire affected the temporal pattern of nectar and pollen available to various pollinator groups, and of fleshy fruits available to all seed-dispersing frugivores. In general, fire seems to have a neutral or even a positive effect on resource availability to mutualists. Nevertheless, there were differences in the availability of the resource utilized by each guild of mutualists.     

Does Argentine ant invasion affect prey availability for foliage-gleaning birds?

Food availability during the breeding season plays a critical role in reproductive success of insectivorous birds. Given that the invasive Argentine ant (Linepithema humile) is known to alter arthropod communities, we predicted that its invasion may affect the availability of food resources for coexisting foliage-gleaning birds. With this aim we studied, for 3 years, foliage arthropods occurring on cork oaks (Quercus suber) and tree heaths (Erica arborea) in invaded and non-invaded secondary forests of the northeastern Iberian Peninsula. Our results show that Argentine ants interact with arboreal foliage arthropods in a different manner than the native ants they displace do. The invasive ant impacted the arthropod community by reducing order diversity and ant species richness and by causing extirpation of most native ant species. Arthropod availability for foliage gleaners’ nestlings diminished in invaded cork oaks, mainly responding to the abundance and biomass depletion of caterpillars. Results suggest that the reproduction of canopy-foraging foliage-gleaning species that mostly rely on caterpillars to feed their young could be compromised by the Argentine ant invasion. Thus, the Argentine ant could be promoting bottom-up effects in the trophic web through its effects on the availability of arthropod preys for insectivorous birds.     

Can temporal covariation and autocorrelation in demographic rates affect population dynamics in a raptor species?

Theoretical studies suggest that temporal covariation among and temporal autocorrelation within demographic rates are important features of population dynamics. Yet, empirical studies have rarely focused on temporal covariation and autocorrelation limiting our understanding of these patterns in natural populations. This lack of knowledge restrains our ability to fully understand population dynamics and to make reliable population forecasts. In order to fill this gap, we used a long‐term monitoring (15 years) of a kestrel Falco tinnunculus population to investigate covariation and autocorrelation in survival and reproduction at the population level and their impact on population dynamics. Using Bayesian joint analyses, we found support for positive covariation between survival and reproduction, but weak autocorrelation through time. This positive covariation was stronger in juveniles compared with adults. As expected for a specialized predator, we found that the reproductive performance was strongly related to an index of vole abundance explaining 86% of the temporal variation. This very strong relationship suggests that the temporally variable prey abundance may drive the positive covariation between survival and reproduction in this kestrel population. Simulations suggested that the observed effect size of covariation could be strong enough to affect population dynamics. More generally, positive covariation and autocorrelation have a destabilizing effect increasing substantially the temporal variability of population size.     

Does energy availability influence classical patterns of spatial variation in exotic species richness?

Aim At macroecological scales, exotic species richness is frequently positively correlated with human population density. Such patterns are typically thought to arise because high human densities are associated with increased introduction effort and/or habitat modification and disturbance. Exotic and native species richness are also frequently positively correlated, although the causal mechanisms remain unclear. Energy availability frequently explains much of the variation in species richness and we test whether such species–energy relationships may influence the relationships of exotic species richness with human population density and native species richness. Location Great Britain. Methods We first investigate how spatial variation in the distributions of the 10 exotic bird species is related to energy availability. We then model exotic species richness using native avian species richness, human population density and energy availability as predictors. Species richness is modelled using two sets of models: one assumes independent errors and the other takes spatial correlation into account. Results The probability of each exotic species occurring, in a 10‐km quadrat, increases with energy availability. Exotic species richness is positively correlated with energy availability, human population density and native species richness in univariate tests. When taking energy availability into account, exotic species richness is negligibly influenced by human population density, but remains positively associated with native species richness. Main conclusions We provide one of the few demonstrations that energy availability exerts a strong positive influence on exotic species richness. Within our data, the positive relationship between exotic species richness and human population density probably arises because both variables increase with energy availability, and may be independent of the influence of human density on the probability of establishment. Positive correlations between exotic and native species richness remain when controlling for the influence of energy on species richness. The relevance of such a finding to the debate on the relationship between diversity and invasibility is discussed.     

Species divergence in offspring begging intensity: difference in need or manipulation of parents?

Conflicts over the delivery and sharing of food among family members are expected to lead to evolution of exaggerated offspring begging for food. Coevolution between offspring begging intensity and parent response depends on the genetic architecture of the traits involved. Given a genetic correlation between offspring begging intensity and parental response, there may be fast and arbitrary divergence in these behaviours between populations. However, there is limited knowledge about the genetic basis of offspring solicitation and parental response and whether these traits are genetically correlated. In this study, we performed a partial cross-fostering experiment of young between pied and collared flycatchers (Ficedula hypoleuca and Ficedula albicollis) and recorded the behaviour of individual offspring and their (foster)parents. We found that nestling collared flycatchers reached a higher phenotypic quality, estimated both as mass at fledging and as intensity of their T-lymphocyte-mediated immune response when raised by heterospecific foster parents. However, although collared flycatchers begged relatively more intensively, we found no evidence of corresponding higher resistance (i.e. lower feeding rate) of adult collared flycatchers than of adult pied flycatchers. Thus, the difference in offspring begging intensity between the two species seems not to be a result of a difference in escalation of the parent-offspring conflict. Instead, the species' divergence in exaggeration of offspring begging intensity 'honestly' matches a difference between the species in offspring need. This interpretation is strengthened by the fact that the difference in begging intensity between the two species increased as the season progressed, coinciding with the higher sensitivity of nestling collared flycatchers to the seasonal decline in food availability. Thus, the behavioural differentiation appears to be a direct consequence of a life-history differentiation (offspring growth patterns).     

Does smoking affect body weight and obesity in China?

An inverse relationship between smoking and body weight has been documented in the medical literature, but the effect of cigarette smoking on obesity remains inconclusive. In addition, the evidence is mixed on whether rising obesity rates are an unintended consequence of successful anti-smoking policies. This study re-examines these relationships using data from China, the largest consumer and manufacturer of tobacco in the world that is also experiencing a steady rise in obesity rates. We focus on the impact of the total number of cigarettes smoked per day on individuals’ body mass index (BMI) and on the likelihood of being overweight and obese. Instrumental variables estimation is used to correct for the endogeneity of cigarette smoking. We find a moderate negative and significant relationship between cigarette smoking and BMI. Smoking is also negatively related to being overweight and obese, but the marginal effects are small and statistically insignificant for being obese. Quantile regression analyses reveal that the association between smoking and BMI is quite weak among subjects whose BMIs are at the high end of the distribution but are considerably stronger among subjects in the healthy weight range. Ordered probit regression analyses also confirm these findings. Our results thus reconcile an inverse average effect of smoking on body weight with the absence of any significant effect on obesity. From a policy perspective these findings suggest that, while smoking cessation may lead to moderate weight gain among subjects of healthy weight, the effects on obese subjects are modest and should not be expected to lead to a large increase in obesity prevalence rates.     

Does carbon enrichment affect hyporheic invertebrates in a gravel stream?

For a period of one year we injected a solution of stream water enriched with glucose and inorganic nitrogen and phosphorus at two experimental sites into the hyporheic sediments of the Oberer Seebach, Austria. The biofilm reacted with a quantitative increase after two weeks. The hyporheic invertebrates were sampled with the Cage Pipe Trap method, where the number of trapped animals is determined by the spatial density and the activity of the invertebrates. Within two and six weeks, the hyporheic invertebrates exhibited a reaction indicating an utilization of the new food resources. Over a longer period of one year, three different reaction patterns appeared. The number of nematods and ostracods increased extensively, presumably caused by the modification of the spatial structure of the environment due to biofilm growth. The number of the small sized invertebrates decreased, reflecting the reduced feeding effort. And the number of the large insect larvae increased indicating that these group is mainly limited by space. The hyporheic zone is described as a ‘self-cleaning DOC filter’, an attribute that is particularly assigned to the ecotone between the riparian soil zone and the stream hyporheic zone.     

Does fluctuating resource availability increase invasibility? Evidence from field experiments in New Zealand short tussock grassland

The theory of fluctuating resource availability proposes that the susceptibility of a plant community to invasion by new species (i.e., invasibility) depends upon conditions of intermittent resource enrichment coinciding with the presence of invading propagules. We compared the response of a rapidly invading forb (Hieracium pilosella L.) between different experimental treatments in a short tussock grassland in New Zealand, over 6–12 years, to determine whether the theory explains differences in invasibility. The theory predicts that environments subject to periodic resource enrichment will be more invasible than those with more stable resource-supply rates. In our study, H. pilosella did not increase more rapidly in treatments subject to periodic resource pulses (fertiliser and water) than in those with more stable resource supplies. Also contrary to the predictions of the theory, the rate of invasion of H. pilosella did not increase following an increase in the rate of supply of water or nutrient resources, or following treatments that temporarily reduced resource uptake in the community, including grazing. H. pilosella did not increase immediately following abrupt increases in water and nutrient supply and removal of the dominant grass species with herbicide, as predicted by the theory, although temporary increases in resident exotic guilds indicated that the intensity of competition for resources was reduced. Neither H. pilosella nor resident exotic guilds showed increased cover growth rates following resumed grazing. The rate of invasion by H. pilosella was not correlated with species richness, a result consistent with one of the predictions of the theory. Therefore, short-lived events that temporarily reduced or suspended competition did not appear to determine the invasion success of this particular species in this region. In New Zealands perennial short tussock grasslands, the characteristics of the resident plant community may be more critical than resource fluctuations in determining invasion success of H. pilosella. Invasion of H. pilosella may be most successfully controlled here by promoting a successional physiognomic shift to a taller, shrub-and-tussock-dominated canopy that competitively excludes low-growing forbs.     

Does thermal history affect metabolic plasticity?: a study in three Phyllotis species along an altitudinal gradient

(1) The aim of this study was to understand the effects of thermal history in metabolic features such as maximum (MMR) and basal (BMR) metabolic rates, as well as in metabolic plasticity, considered as the total variation of MMR and BMR during the acclimation period. (2) We studied three species of the genus Phyllotis, from different thermal environments, in an altitudinal gradient from sea level to 3800m.a.s.l. Animals were acclimated to contrasting temperatures of 5 and 30 degrees C. To determine the metabolic flexibility, MMR was measured at intervals of 6 days during the acclimation period, while BMR values were obtained at the end of acclimations. Aerobic scope and the rates of change of MMR were estimated in all populations. (3) High- and low-altitude rodents did not show differences in BMR. However, both upper and lower limits of MMR, as well as aerobic scope, were significantly different between high- and low-altitude species, indicating similar ranges of metabolic plasticity. On the other hand, the rates of change of MMR were similar in all populations. (4) Our results indicate that thermal history has a profound effect on the individuals' thermogenic capacity, probably in both phylogenetic and ontogenetic levels. Low-altitude species could not increase MMR to the same levels as high-altitude species, while the later were unable to decrease MMR to achieve the values of the low-altitude species.