Adult survival probability and body size affect parental risk-taking across latitudes.

Parents faced with a predator must choose between their own safety versus taking care of their offspring. Each choice can have fitness costs. Life-history theory predicts that longer-lived species should be less willing than shorter-lived species to return to care for their offspring after a predator disturbance because they have more opportunities to reproduce in the future. We increased adult predation risk during incubation for 40 bird species in north temperate, tropical, and south temperate latitudes. We found that species with higher adult survival probabilities were more cautious, waiting longer before returning to the nest to provide care. Contrary to other studies, we also found that parents were more risk averse and waited longer to return in smaller than larger species, likely reflecting greater vulnerability of smaller species. Ultimately, the relative risk a predator poses to a species and the probability of future reproduction predict parental risk taking across the world.

Do avian species survive better on islands?

Island species are often predictably different from their mainland counterparts. Milder climates and reduced predation risk on islands have been involved to explain shifts in body size and a suite of life-history traits such as clutch size and offspring growth rate. Despite the key role of adult survival on risk taking and reproduction, the prediction that living on islands increases adult survival has yet to be tested systematically. I gathered data on adult annual apparent survival from the island and mainland year-round resident species of birds from around the world. With this large dataset (697 species), I found that species of birds living on islands showed higher apparent survival than their mainland counterparts in the two Hemispheres and at all latitudes, controlling for several known predictors of adult survival, including body size, clutch size and breeding system. These results shed light on the ecological factors that influence survival on islands and extend the life-history island syndrome to adult survival.

Flocking in birds increases annual adult survival in a global analysis.

Adaptive hypotheses for the evolution of group foraging in animals typically invoke enhanced foraging efficiency and reduced predation risk. Net benefits of group foraging in different species should translate into a survival advantage for group members. Despite numerous interspecific studies in birds and mammals, few have documented a survival advantage for group foraging species. Using a large dataset in birds (> 1100 species worldwide), I investigated whether annual adult apparent survival was higher in species that forage in flocks than in solitary species. Using a phylogenetic framework to account for relatedness among species and controlling for known correlates of adult survival in birds such as body size, clutch size, latitude, and diet, I documented a positive effect of flocking on annual adult apparent survival. The increase in survival was less pronounced in occasionally flocking species suggesting that the benefits of group foraging can depend on the frequency of its use. The results highlight how group foraging can increase fitness in animals.

Exposure to essential oils and ethanol vapors affect fecundity and survival of two frugivorous fruit fly (Diptera: Tephritidae) pest species.

Plant-derived compounds can be an environmentally friendly alternative to synthetic pesticide use for pest management. Essential oils (EOs) in several plant families have been found to be toxic to various pest species of insects through topical application, ingestion, and as fumigants. Previous studies revealed that, among various environmentally friendly insecticides, the EOs of Baccharis dracunculifolia and Pinus elliottii and an ethanol extract of Solanum granulosoleprosum plus Ricinus communis, were toxic to Ceratitis capitata and Anastrepha fraterculus (Diptera: Tephritidae) when applied topically to pupae or when ingested by adults. Here, we aimed to examine the potentially toxic effects of these plant-derived compounds when these two pestiferous fruit fly species were exposed to their vapors. We also examined their fumigant effect on female fecundity and fertility and compared it with water and ethanol controls. Exposure of C. capitata and A. fraterculus sexually mature adults to volatiles and vapors of both B. dracunculifolia and P. elliottii EOs resulted in lower longevity (half-life), survivorship, and female fecundity than the water vapor control. Toxicity of C. capitata was greater for P. elliottii than for B. dracunculifolia while the reverse was true for A. fraterculus. Exposure to vapors of S. granulosoleprosum + R. communis (S + R) had no effect on longevity but reduced survivorship of adults of both species. Interestingly, exposure to vapors of S + R, 50% (v/v) and pure ethanol resulted in greater fecundity of females of both frugivorous fly species than the water control. By contrast, fertility (% egg hatch) was in all cases high (>85%) and not different than the water control. Exposure to ethanol vapors appears to have similar effects on frugivorous tephritids as those reported on saprophagous and frugivorous species of Drosophila, a novel finding that may have important practical implications.

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season.

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long-lived seabird that experiences high bycatch rates in longline fisheries and strong population-level impacts due to this type of anthropogenic mortality. Analyzing a long-term dataset on individual monitoring, we compared adult survival (by means of multi-event capture-recapture models) among three close predator-free Mediterranean colonies of the species. Unexpectedly for a long-lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony-specific and mortality risk varies spatially.

Resource Allocation and Seed Size Selection in Perennial Plants under Pollen Limitation.

Pollen limitation may affect resource allocation patterns in plants, but its role in the selection of seed size is not known. Using an evolutionarily stable strategy model of resource allocation in perennial iteroparous plants, we show that under density-independent population growth, pollen limitation (i.e., a reduction in ovule fertilization rate) should increase the optimal seed size. At any level of pollen limitation (including none), the optimal seed size maximizes the ratio of juvenile survival rate to the resource investment needed to produce one seed (including both ovule production and seed provisioning); that is, the optimum maximizes the fitness effect per unit cost. Seed investment may affect allocation to postbreeding adult survival. In our model, pollen limitation increases individual seed size but decreases overall reproductive allocation, so that pollen limitation should also increase the optimal allocation to postbreeding adult survival. Under density-dependent population growth, the optimal seed size is inversely proportional to ovule fertilization rate. However, pollen limitation does not affect the optimal allocation to postbreeding adult survival and ovule production. These results highlight the importance of allocation trade-offs in the effect pollen limitation has on the ecology and evolution of seed size and postbreeding adult survival in perennial plants.

Contrasting effects of tropical cyclones on the annual survival of a pelagic seabird in the Indian Ocean.

Tropical cyclones are renowned for their destructive nature and are an important feature of marine and coastal tropical ecosystems. Over the last 40 years, their intensity, frequency and tracks have changed, partly in response to ocean warming, and future predictions indicate that these trends are likely to continue with potential consequences for human populations and coastal ecosystems. However, our understanding of how tropical cyclones currently affect marine biodiversity, and pelagic species in particular, is limited. For seabirds, the impacts of cyclones are known to be detrimental at breeding colonies, but impacts on the annual survival of pelagic adults and juveniles remain largely unexplored and no study has simultaneously explored the direct impacts of cyclones on different life-history stages across the annual life cycle. We used a 20-year data set on tropical cyclones in the Indian Ocean, tracking data from 122 Round Island petrels and long-term capture-mark-recapture data to explore the impacts of tropical cyclones on the survival of adult and juvenile (first year) petrels during both the breeding and migration periods. The tracking data showed that juvenile and adult Round Island petrels utilize the three cyclone regions of the Indian Ocean and were potentially exposed to cyclones for a substantial part of their annual cycle. However, only juvenile petrel survival was affected by cyclone activity; negatively by a strong cyclone in the vicinity of the breeding colony and positively by increasing cyclone activity in the Northern Indian Ocean where they spend the majority of their first year at sea. These contrasting effects raise the intriguing prospect that the projected changes in cyclones under current climate change scenarios may have positive as well as the more commonly perceived negative impacts on marine biodiversity.

Surviving at high elevations: an inter- and intra-specific analysis in a mountain bird community.

Elevation represents an important selection agent on self-maintenance traits and correlated life histories in birds, but no study has analysed whether life-history variation along this environmental cline is consistent among and within species. In a sympatric community of passerines, we analysed how the average adult survival of 25 open-habitat species varied with their elevational distribution and how adult survival varied with elevation at the intra-specific level. For such purpose, we estimated intra-specific variation in adult survival in two mountainous species, the Water pipit (Anthus spinoletta) and the Northern wheatear (Oenanthe oenanthe) in NW Spain, by means of capture-recapture analyses. At the inter-specific level, high-elevation species showed higher survival values than low elevation ones, likely because a greater allocation to self-maintenance permits species to persist in alpine environments. At the intra-specific level, the magnitude of survival variation was lower by far. Nevertheless, Water pipit survival slightly decreased at high elevations, while the proportion of transient birds increased. In contrast, no such relationships were found in the Northern wheatear. Intra-specific analyses suggest that living at high elevation may be costly, such as for the Water pipit in our case study. Therefore, it seems that a species can persist with viable populations in uplands, where extrinsic mortality is high, by increasing the investment in self-maintenance and prospecting behaviours.

Factors that influence vital rates of Seaside and Saltmarsh sparrows in coastal New Jersey, USA.

As saltmarsh habitat continues to disappear, understanding the factors that influence saltmarsh breeding bird population dynamics is an important step for the conservation of these declining species. Using five years (2011 - 2015) of demographic data, we evaluated and compared Seaside (Ammodramus maritimus) and Saltmarsh (A. caudacutus) sparrow apparent adult survival and nest survival at the Edwin B. Forsythe National Wildlife Refuge, New Jersey, USA. We determined the effect of site management history (unditched vs. ditched marsh) on adult and nest survival to aid in prioritizing future management or restoration actions. Seaside Sparrow apparent adult survival (61.6%, 95% CI: 52.5 - 70.0%) averaged >1.5 times greater than Saltmarsh Sparrow apparent adult survival (39.9%, 95% CI: 34.0 - 46.2%). Nest survival and predation and flooding rates did not differ between species, and predation was the primary cause of failure for both species. Apparent adult survival and nest survival did not differ between unditched and ditched marshes for either species, indicating that marsh ditching history may not affect breeding habitat quality for these species. With predation as the primary cause of nest failure for both species in New Jersey, we suggest that future research should focus on identification of predator communities in salt marshes and the potential for implementing predator-control programs to limit population declines.

Opposing selection and environmental variation modify optimal timing of breeding.

Studies of evolution in wild populations often find that the heritable phenotypic traits of individuals producing the most offspring do not increase proportionally in the population. This paradox may arise when phenotypic traits influence both fecundity and viability and when there is a tradeoff between these fitness components, leading to opposing selection. Such tradeoffs are the foundation of life history theory, but they are rarely investigated in selection studies. Timing of breeding is a classic example of a heritable trait under directional selection that does not result in an evolutionary response. Using a 22-y study of a tropical parrot, we show that opposing viability and fecundity selection on the timing of breeding is common and affects optimal breeding date, defined by maximization of fitness. After accounting for sampling error, the directions of viability (positive) and fecundity (negative) selection were consistent, but the magnitude of selection fluctuated among years. Environmental conditions (rainfall and breeding density) primarily and breeding experience secondarily modified selection, shifting optimal timing among individuals and years. In contrast to other studies, viability selection was as strong as fecundity selection, late-born juveniles had greater survival than early-born juveniles, and breeding later in the year increased fitness under opposing selection. Our findings provide support for life history tradeoffs influencing selection on phenotypic traits, highlight the need to unify selection and life history theory, and illustrate the importance of monitoring survival as well as reproduction for understanding phenological responses to climate change.

Some observations on overwintering sites of adult Culex quinquefasciatus (Diptera: Culicidae) and strategies followed under natural and seminatural conditions.

Field population of adult Culex quinquefasciatus Say, landed in December, congregated and overwintered in indoor artificial sites in Delhi. Repeated sampling strategy by individually collecting the adults was adopted to study their overwintering strategies for 4 years (December to April). They remained vagile and readily repopulated the resting sites after the removal of samples. A large percentage of females was fertile, unfed and nulliparous indicating that reproduction ceased in them. Adult survival was significantly prolonged to a maximum of 3 months under natural conditions. Gonotrophic cycle also got prolonged. Close to quitting, they became gravid and left in April without oviposition. No adults were observed on the sites for the rest of the year. Oviposition was induced in the blood-engorged females when provided with food, water and outdoor conditions. Oviposition might have been induced directly by water and food provided them energy under seminatural conditions. Eggs were laid singly or in the form of rafts, and the number in both the cases was low. Singly laid eggs did not hatch, and in rafts, hatching was ~80 %. Winter conditions seemed to strongly impact fertility, blood feeding, fecundity, oviposition behaviour, egg hatchability and longevity. Use of the overwintering sites as biological tool, as a part of environmental control in IPM, is suggested for organising antivector measures during winter. There is a need of exploring and creating more sites of this kind.

Interspecific Larval Competition Differentially Impacts Adult Survival in Dengue Vectors.

Mosquitoes often experience intraspecific and interspecific competition among larvae attributable to high densities and nutrient limitation, especially container mosquitoes including Aedes aegypti (L.) and Aedes albopictus (Skuse). Density-dependent effects on larvae impact adult production and adult traits that influence transmission of arboviruses. To improve our understanding of the mechanisms by which density-dependence influences transmission and identify species-specific traits, we tested the hypotheses: 1) Competitive asymmetry in favor of Ae. albopictus over Ae. aegypti translates to altered adult female survival, and 2) Ae. aegypti adult females are more resistant to life-shortening effects of low-humidity conditions than Ae. albopictus. We gauged the relative impact of inter- and intraspecific larval competition on adult survival in high- and low-humidity regimes (77 and 44% relative humidity, respectively). For Ae. albopictus, intraspecific but not interspecific competition usually reduced adult survival under both humidity regimes. For Ae. aegypti, both intraspecific and interspecific competition reduced adult survival. Ae. albopictus adult survival was minimally influenced by interspecific competition with Ae. aegypti, consistent with observations that Ae. albopictus is the superior competitor. A species comparison indicated that Ae. aegypti exhibited a survival advantage relative to Ae. albopictus under both low- and high-humidity conditions. However, similar survival of these Aedes species was observed in some cases depending on conditions experienced in both the aquatic and terrestrial environments. These results demonstrate plasticity in survival rates of dengue and chikungunya vectors and the significance of considering the influence of biological interactions during the immature stages and abiotic conditions during the adult stage.

Evaluation of Bacillus thuringiensis israelensis as a Control Agent for Adult Anopheles gambiae.

Unlike the application of Bacillus thuringiensis israelensis (Bti) for the control of larval mosquitoes, studies of its effects on adults, for its possible use in attractive toxic sugar baits, have resulted in conflicting results. Five species have shown a decrease in adult survival due to Bti ingestion, whereas adults of Anopheles arabiensis have not. We sought to determine if ingestion of Bti by adults of Anopheles gambiae, a sibling species of An. arabiensis, increases their mortality. Laboratory-reared adults were provided continuously from emergence with water only, a sucrose solution, or a Bti suspension in sucrose solution. After 3 days, the Bti suspension was replaced with untainted sucrose solution. The mosquitoes with only water were all dead by day 3. The survivorships of those in the sucrose and sucrose-Bti treatments were insignificantly different, both with an LT50 (Lethal Time, time until 50% of individuals died) of 25 days. The results support the conclusion that adult survivorship of An. gambiae-complex members is unaffected by the ingestion of Bti in sugar meals.

Synchrony in adult survival is remarkably strong among common temperate songbirds across France.

Synchronous variation in demographic parameters across species increases the risk of simultaneous local extinction, which lowers the probability of subsequent recolonization. Synchrony therefore tends to destabilize meta-populations and meta-communities. Quantifying interspecific synchrony in demographic parameters, like abundance, survival, or reproduction, is thus a way to indirectly assess the stability of meta-populations and meta-communities. Moreover, it is particularly informative to identify environmental drivers of interspecific synchrony because those drivers are important across species. Using a Bayesian hierarchical multisite multispecies mark-recapture model, we investigated temporal interspecific synchrony in annual adult apparent survival for 16 common songbird species across France for the period 2001-2016. Annual adult survival was largely synchronous among species (73%, 95% credible interval [47%-94%] of the variation among years was common to all species), despite species differing in ecological niche and life history. This result was robust to different model formulations, uneven species sample sizes, and removing the long-term trend in survival. Synchrony was also shared across migratory strategies, which suggests that environmental forcing during the 4-month temperate breeding season has a large-scale, interspecific impact on songbird survival. However, the strong interspecific synchrony was not easily explained by a set of candidate weather variables we defined a priori. Spring weather variables explained only 1.4% [0.01%-5.5%] of synchrony, while the contribution of large-scale winter weather indices may have been stronger but uncertain, accounting for 12% [0.3%-37%] of synchrony. Future research could jointly model interspecific variation and covariation in breeding success, age-dependent survival, and age-dependent dispersal to understand when interspecific synchrony in abundance emerges and destabilizes meta-communities.

Cascading effects of pre-adult survival on sexual selection.

Sexual selection influences broad-scale patterns of biodiversity. While a large body of research has investigated the effect of mate competition on sexual selection, less work has examined how pre-adult life history influences sexual selection. We used a mathematical framework to explore the influence of pre-adult survival on sexual selection. Our model suggests that pre-adult male mortality will affect the strength of sexual selection when a fixed number of adult males have an advantageous mate-acquisition trait. When a fixed number of males have an advantageous mate-acquisition trait, sexual selection is expected to increase when pre-adult mortality is relatively low. By contrast, if a fixed proportion (rather than number) of adult males have a mate-acquisition trait, pre-adult male mortality is not expected to affect the strength of sexual selection. Further, if the advantageous mating trait affects pre-adult survival, natural and sexual selection can interact to influence the overall selection on the mating trait. Given that pre-adult mortality is often shaped by natural selection, our results highlight conditions under which natural selection can have cascading effects on sexual selection.

Comparison of Population Aging in Europe and Asia Using a Time-Consistent and Comparative Aging Measure.

Objective: We compare population aging in Europe and Asia using a measure that is both consistent over time and appropriate for cross-country comparison. Method: Sanderson and Scherbov proposed to estimate the old-age threshold by the age at which the remaining life expectancy (RLE) equals 15 years. We propose an adjustment of this measure, taking into account cross-national differences in the exceptionality of reaching that age. Results: Our old-age threshold was lower than 65 years in 2012 in Central Asia, Southern Asia, Southeastern Asia, and many Eastern European countries. These populations also experienced a higher share of elderly compared with the RLE15 method. Our method revealed more geographical diversity in the shares of elderly. Both methods exhibited similar time trends for the old-age thresholds and the shares of elderly. Discussion: Our prospective and comparative measure reveals higher population aging estimates in most Asian and Eastern European countries and more diversity in aging.

The Survival of Mediterranean Fruit Fly (Diptera: Tephritidae) Over Winter in Western Australia.

The Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) is one of the most economically important pest insects of fruit crops worldwide. Mediterranean fruit fly can cause up to 100% crop loss in susceptible fruit. In order to formulate best management practices, it is critical to understand how Mediterranean fruit fly overwinters in a given geographical location and bridge the gap between autumn and spring populations. In this study, we evaluated the overwintering potential of Mediterranean fruit fly immature and adult stages in two locations in Perth Hills, Western Australia. We also monitored wild adult Mediterranean fruit fly populations for 2 yr. Adults were present year-round with captures very low in winter to early spring relative to summer and autumn. Field experiments revealed that immature stages in apples (eggs/first instar) and soil (pupae) remained viable in winter, emerging as adults at the onset of warmer weather in spring. In field cages, adults survived 72-110 d, and female laid viable eggs when offered citrus fruit, though only 1-6% eggs survived to emerge as adults. Adults survived longer in field cages when offered live citrus branch. The findings suggest that all Mediterranean fruit fly life stages can survive through mild winter, and surviving adults, eggs in the fruit and/or pupae in the soil are the sources of new population that affect the deciduous fruit crops in Perth. We recommend that Mediterranean fruit fly monitoring is required year-round and control strategies be deployed in spring. Furthermore, we recommend removal of fallen fruit particularly apple and other winter fruit such as citrus.

Reproduction and survival in the city: which fitness components drive urban colonization in a reed-nesting waterbird?

Processes of adaptation to urban environments are well described for relatively few avian taxa, mainly passerines, but selective forces responsible for urban colonization in ecologically different groups of birds remain mostly unrecognized. The aim of this article is to identify drivers of recent urban colonization (Lódz, central Poland) by a reed-nesting waterbird, the Eurasian coot Fulica atra. Urban colonizers were found to adopt a distinct reproductive strategy by maximizing the number of offspring (carryover effects of higher clutch size), whereas suburban individuals invested more in the quality of the progeny (higher egg volume), which could reflect differences in predatory pressure between 2 habitats. In fact, reduced predation rate was strongly suggested by elevated hatching success in highly urbanized areas, where probability of hatching at least 1 chick was higher by 30% than in suburban natural-like habitats. Coots nesting in highly urbanized landscape had considerably higher annual reproductive success in comparison to suburban pairs, and the difference was 4-fold between the most and least urbanized areas. There was also a constant increase in size-adjusted body mass and hemoglobin concentration of breeding coots from the suburbs to the city centre. Urban colonization yielded no survival benefits for adult birds and urban individuals showed higher site fidelity than suburban conspecifics. The results suggest that the recent urban colonization by Eurasian coots was primary driven by considerable reproductive benefits which may be primarily attributed to: (1) reduced predation resulting from an exclusion of most native predators from highly urbanized zones; (2) increased condition of urban-dwelling birds resulting from enhanced food availability.

Adult Survival and Imperfect Assessment of Parentage: Effects on Male Parenting Decisions.

Applications of molecular methods to assess parentage have revealed that the distribution of reproductive success among individuals often differs, sometimes dramatically, from expectation based on observation of behavioral association. Much theory exists on whether and when males should reduce parental care in response to level of paternity. Life-history theory predicts that trade-offs in reproductive effort should be influenced by adult survival. We used a dynamic programming approach to address how level of paternity, ability to assess paternity, and adult survival rate interact to affect male tolerance of reduced parentage in a given brood. Adult survival has the greatest influence on male decisions such that, for any given cost of reproduction and value of male care, tolerance of extrapair fertilizations (EPFs) decreases as adult survival increases. An unexpected result of these models is that an optimal response also depends on a male's ability to predict probability of parentage (i.e., uncertainty). These models better characterize the nature of paternity uncertainty and its effect on EPF tolerance than have previous models and add to our understanding of the complex relationship between uncertainty, mating strategies, and adult survival.

Food availability and nest predation influence life history traits in Audouin's gull, Larus audouinii.

The effects of food availability and nest predation on several life history traits such as adult survival, dispersal, and reproductive performance were assessed in an Audouin's gull (Larus audouinii) colony during the period 1992-1997. The amounts of fish discarded from trawlers were used as a measure of food availability, and a trawling moratorium which partially overlapped with the breeding season of the gulls was taken into account. The effects of nest predation were assessed in 1994, when a terrestrial predator entered the colony and remained for the whole breeding season preying on both eggs and chicks. Using the moratorium and the predatory event as natural experiments, several hypotheses were tested: (a) food supply would affect breeding performance but not adult survival (independently of age and sex), since gulls are long-lived and adult survival is the most sensitive demographic parameter in their population dynamics; (b) the predator would trigger breeding dispersal (although gulls are mostly philopatric, they are known to abandon their natal colony after breeding failure instigated by events such as this). If breeding dispersal occurs, the rate is expected to be higher in females than in males, and higher in new breeders than in more experienced breeding birds, as is usually recorded in colonial seabirds. Probabilities of resighting and survival were estimated separately, using capture-recapture models. As expected, changes in food availability did not affect adult survival, whereas they influenced egg volume, clutch size, and breeding success. Local adult survival was estimated to be 0.908 (SD = 0.007) for males and females, and it did not change significantly with the age of individuals (range 3-8 years). The predator significantly decreased breeding success, and caused the dispersal of a number of adults probably to breed in another colony; this rate was estimated at an average of 0.10 (SD = 0.02). As expected, inexperienced breeders dispersed significantly more (14%) than more experienced breeders (8%) after the predator event, but dispersal was not sex biased. Recapture probabilities after the predator event suggest that birds that left the colony still had not returned. Results confirm that population dynamics of ground-nesting seabirds are sensitive to terrestrial predation, even when predation caused only a partial breeding failure.