Ocean climate and seal condition

Background  

The condition of many marine mammals varies with fluctuations in productivity and food supply in the ocean basin where they forage. Prey is impacted by physical environmental variables such as cyclic warming trends. The weaning weight of northern elephant seal pups, Mirounga angustirostris, being closely linked to maternal condition, indirectly reflects prey availability and foraging success of pregnant females in deep waters of the northeastern Pacific. The aim of this study was to examine the effect of ocean climate on foraging success in this deep-diving marine mammal over the course of three decades, using cohort weaning weight as the principal metric of successful resource accrual.     

Climate influences on female survival in a declining population of southern elephant seals (Mirounga leonina)

The Southern Ocean has been disproportionately affected by climate change and is therefore an ideal place to study the influence of changing environmental conditions on ecosystems. Changes in the demography of predator populations are indicators of broader shifts in food web structure, but long‐term data are required to study these effects. Southern elephant seals (Mirounga leonina) from Macquarie Island have consistently decreased in population size while all other major populations across the Southern Ocean have recently stabilized or are increasing. Two long‐term mark‐recapture studies (1956–1967 and 1993–2009) have monitored this population, which provides an opportunity to investigate demographic performance over a range of climatic conditions. Using a 9‐state matrix population model, we estimated climate influences on female survival by incorporating two major climatic indices into our model: The Southern Annular Mode (SAM) and the Southern Oscillation Index (SOI). Our best model included a 1 year lagged effect of SAM and an unlagged SOI as covariates. A positive relationship with SAM1 (lagged) related the previous year''s SAM with juvenile survival, potentially due to changes in local prey availability surrounding Macquarie Island. The unlagged SOI had a negative effect on both juvenile and adult seals, indicating that sea ice dynamics and access to foraging grounds on the East Antarctic continental shelf could explain the different contributions of ENSO events on the survival of females in this population.     

Effects of large-scale climatic fluctuations on survival and production of young in a Neotropical migrant songbird, the yellow warbler Dendroica petechia

Migrant songbirds are vulnerable to changes in climatic conditions on both the breeding and wintering grounds. For North American Neotropical migrants, the El Niño/Southern Oscillation (ENSO), via its effects on global precipitation and temperature, modulates the productivity of their temperate and tropical terrestrial ecosystems used during the course of their annual cycle. We evaluated how a densely nesting population of yellow warblers Dendroica petechia in a riparian forest in southern Manitoba, Canada, responded to the El Niño/Southern Oscillation (ENSO) between 1992 and 2001. Standardized mist netting was used to estimate apparent annual survival of adults and production of young. Both adult survival and the production of young were positively correlated with the Southern Oscillation Index (SOI). More specifically, values of both these demographic parameters were lowest during El Niño years and highest during La Niña years. These findings demonstrate the influence of climate on populations of Neotropical migrants in North America. The more frequent El Niños predicted to result from future global climate change could negatively affect populations of yellow warblers and other Neotropical migrants breeding in this region.     

The timing of life-history events in a changing climate

Although empirical and theoretical studies suggest that climate influences the timing of life-history events in animals and plants, correlations between climate and the timing of events such as egg-laying, migration or flowering do not reveal the mechanisms by which natural selection operates on life-history events. We present a general autoregressive model of the timing of life-history events in relation to variation in global climate that, like autoregressive models of population dynamics, allows for a more mechanistic understanding of the roles of climate, resources and competition. We applied the model to data on 50 years of annual dates of first flowering by three species of plants in 26 populations covering 4 degrees of latitude in Norway. In agreement with earlier studies, plants in most populations and all three species bloomed earlier following warmer winters. Moreover, our model revealed that earlier blooming reflected increasing influences of resources and density-dependent population limitation under climatic warming. The insights available from the application of this model to phenological data in other taxa will contribute to our understanding of the roles of endogenous versus exogenous processes in the evolution of the timing of life-history events in a changing climate.     

Relative costs of offspring sex and offspring survival in a polygynous mammal

Costs of reproduction are expected to be ubiquitous in wild animal populations and understanding the drivers of variation in these costs is an important aspect of life-history evolution theory. We use a 43 year dataset from a wild population of red deer to examine the relative importance of two factors that influence the costs of reproduction to mothers, and to test whether these costs vary with changing ecological conditions. Like previous studies, our analyses indicate fitness costs of lactation: mothers whose calves survived the summer subsequently showed lower survival and fecundity than those whose calves died soon after birth, accounting for 5% and 14% of the variation in mothers'' survival and fecundity, respectively. The production of a male calf depressed maternal survival and fecundity more than production of a female, but accounted for less than 1% of the variation in either fitness component. There was no evidence for any change in the effect of calf survival or sex with increasing population density.     

The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long-lived seabird

1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.     

Weather-dependent survival: implications of climate change for passerine population processes

Understanding demographic processes will be essential to construct robust models of population responses to climate change. We show that survival is related to the strength of the North Atlantic Oscillation in five out of ten British resident passerine species, and explore the importance of biologically more specific variables (duration of winter frosts and snow periods; occurrence of cold, wet days; spring temperature; and summer drought). The most important variables differed between species in relation to differences in foraging strategy. In almost all cases, first-year survival was influenced by weather more than was the survival of adult birds. Particularly vulnerable species, such as the Wren Troglodytes troglodytes , may exhibit a 25% reduction in juvenile survival rates due to adverse weather within the range experienced in the last 30 years; variation in survival by 10% or more is commonplace in most species. Thus, climate influences on food availability may provide the mechanism by which populations will alter under changed climatic conditions, though the presence of density dependence may reduce the impact of this on long-term population trajectories.     

Environmental forcing and Southern Ocean marine predator populations: effects of climate change and variability

The Southern Ocean is a major component within the global ocean and climate system and potentially the location where the most rapid climate change is most likely to happen, particularly in the high-latitude polar regions. In these regions, even small temperature changes can potentially lead to major environmental perturbations. Climate change is likely to be regional and may be expressed in various ways, including alterations to climate and weather patterns across a variety of time-scales that include changes to the long interdecadal background signals such as the development of the El Niño–Southern Oscillation (ENSO). Oscillating climate signals such as ENSO potentially provide a unique opportunity to explore how biological communities respond to change. This approach is based on the premise that biological responses to shorter-term sub-decadal climate variability signals are potentially the best predictor of biological responses over longer time-scales. Around the Southern Ocean, marine predator populations show periodicity in breeding performance and productivity, with relationships with the environment driven by physical forcing from the ENSO region in the Pacific. Wherever examined, these relationships are congruent with mid-trophic-level processes that are also correlated with environmental variability. The short-term changes to ecosystem structure and function observed during ENSO events herald potential long-term changes that may ensue following regional climate change. For example, in the South Atlantic, failure of Antarctic krill recruitment will inevitably foreshadow recruitment failures in a range of higher trophic-level marine predators. Where predator species are not able to accommodate by switching to other prey species, population-level changes will follow. The Southern Ocean, though oceanographically interconnected, is not a single ecosystem and different areas are dominated by different food webs. Where species occupy different positions in different regional food webs, there is the potential to make predictions about future change scenarios.     

Divergent patterns of impact of environmental conditions on life history traits in two populations of a long-distance migratory bird

Some areas have experienced recent dramatic warming due to climate change, while others have shown no change at all, or even recent cooling. We predicted that patterns of selection on life history would differ between southern and northern European populations of a long-distance migratory bird, the barn swallow Hirundo rustica, because global patterns of weather as reflected by large-scale weather phenomena such as the North Atlantic Oscillation (NAO) and the El Niño-Southern Oscillation (ENSO) have different effects on environmental conditions in different parts of the world frequented during the annual cycle. We investigated relationships between mean arrival date, dispersal rate and yearling survival rate among years, using two long-term population studies in Spain and Denmark. We found evidence of a difference in the effects of normalized difference vegetation index in North and West Africa on mean arrival date of male barn swallows, with the effect differing significantly between populations. Second, there was a significant interaction between ENSO and population on dispersal rate, showing that conditions in Africa during winter differentially affected dispersal in the two populations. Finally, the NAO index in winter had an effect on yearling survival that differed between populations. These findings highlight the divergent patterns of response to climate change among populations, and they suggest that climate change can differentially affect important life history traits with potential implications for maintenance of viable populations and gene flow among populations.     

Climate and body size influence nest survival in a fish with parental care

1. The current study examined the effect of broad-scale climate and individual-specific covariates on nest survival in smallmouth bass over a 20-year period. 2. Large-scale climate indices [winter North Atlantic Oscillation (NAO) and winter El Ni?o/Southern Oscillation (ENSO)] and body size of parental males were important covariates in nest survival along with nest age and a quadratic trend in survival. 3. We did not find an effect due to a habitat covariate (total effective fetch) or a phenology covariate (degree-days at start of nesting) on nest survival. 4. Male size in the second half of the nesting season was a more influential covariate on nest success than male size in the first half or throughout the nesting period. 5. We present evidence showing that winter NAO/ENSO indices establish limnological conditions the following spring that influence thermal stability of the lake during the nesting period. 6. The combined climate and body size covariates point to nest survival as a function of lagged climate-scale influences on limnology and the individual-scale influence of bioenergetics on the duration of parental care and nest success.     

Hormonally mediated maternal effects, individual strategy and global change

A challenge to ecologists and evolutionary biologists is predicting organismal responses to the anticipated changes to global ecosystems through climate change. Most evidence suggests that short-term global change may involve increasing occurrences of extreme events, therefore the immediate response of individuals will be determined by physiological capacities and life-history adaptations to cope with extreme environmental conditions. Here, we consider the role of hormones and maternal effects in determining the persistence of species in altered environments. Hormones, specifically steroids, are critical for patterning the behaviour and morphology of parents and their offspring. Hence, steroids have a pervasive influence on multiple aspects of the offspring phenotype over its lifespan. Stress hormones, e.g. glucocorticoids, modulate and perturb phenotypes both early in development and later into adulthood. Females exposed to abiotic stressors during reproduction may alter the phenotypes by manipulation of hormones to the embryos. Thus, hormone-mediated maternal effects, which generate phenotypic plasticity, may be one avenue for coping with global change. Variation in exposure to hormones during development influences both the propensity to disperse, which alters metapopulation dynamics, and population dynamics, by affecting either recruitment to the population or subsequent life-history characteristics of the offspring. We suggest that hormones may be an informative index to the potential for populations to adapt to changing environments.     

Legacies of La Niña: North American monsoon can rescue trees from winter drought

While we often assume tree growth–climate relationships are time‐invariant, impacts of climate phenomena such as the El Niño Southern Oscillation (ENSO) and the North American Monsoon (NAM) may challenge this assumption. To test this assumption, we grouped ring widths (1900‐present) in three southwestern US conifers into La Niña periods (LNP) and other years (OY). The 4 years following each La Niña year are included in LNP, and despite 1–2 year growth declines, compensatory adjustments in tree growth responses result in essentially equal mean growth in LNP and OY, as average growth exceeds OY means 2–4 years after La Niña events. We found this arises because growth responses in the two periods are not interchangeable: Due to differences in growth–climate sensitivities and climatic memory, parameters representing LNP growth fail to predict OY growth and vice versa (decreases in R² up to 0.63; lowest R² = 0.06). Temporal relationships between growth and antecedent climate (memory) show warmer springs and longer growing seasons negatively impact growth following dry La Niña winters, but that NAM moisture can rescue trees after these events. Increased importance of monsoonal precipitation during LNP is key, as the largest La Niña‐related precipitation deficits and monsoonal precipitation contributions both occur in the southern part of the region. Decreases in first order autocorrelation during LNP were largest in the heart of the monsoon region, reflecting both the greatest initial growth declines and the largest recovery. Understanding the unique climatic controls on growth in Southwest conifers requires consideration of both the influences and interactions of drought, ENSO, and NAM, each of which is likely to change with continued warming. While plasticity of growth sensitivity and memory has allowed relatively quick recovery in the tree‐ring record, recent widespread mortality events suggest conditions may soon exceed the capacity for adjustment in current populations.     

Radiocollars do not affect the survival or foraging behaviour of wild meerkats

The practice of radiocollaring is increasingly being used in the study of wild animal populations. However, methods of tracking and monitoring are known to affect some species negatively. On a wild population of habituated meerkats Suricata suricatta , we investigated the potential costs and consequences of carrying a radiocollar for small carnivores, focusing on detailed aspects of foraging behaviour and survival. Radiocollared individuals were no more likely to be predated than non-collared individuals in the same social group. We also found that collaring did not affect foraging efficiency, with collared individuals being equally efficient foragers (four measures) while wearing a collar compared with the pre-collaring period. To control for group-specific events, changes in foraging efficiency after collaring were compared with the behaviour of control individuals from the same group and no significant differences were found. In conclusion, we found no evidence to suggest radiocollars impact negatively on the bearer's welfare, survival or foraging ability.     

DO EMBRYOS INFLUENCE MATERNAL INVESTMENT? EVALUATING MATERNAL‐FETAL COADAPTATION AND THE POTENTIAL FOR PARENT‐OFFSPRING CONFLICT IN A PLACENTAL FISH

The evolution of matrotrophy introduces the potential for genomic conflicts between mothers and embryos. These conflicts are hypothesized to accelerate the evolution of reproductive isolation and to influence the evolution of life-history traits, reproductive structures, and genomic imprinting. These hypotheses assume offspring can influence the amount of maternal investment they receive and that there is a trade-off between maternal investment into individual offspring and maternal survival or fecundity. We used field data and laboratory crosses to test whether these assumptions are met in the matrotrophic poeciliid fish Heterandria formosa . Comparisons of life histories between two natural populations demonstrated a trade-off between the level of maternal investment into individual embryos and maternal fecundity. Laboratory crosses between individuals from these populations revealed that offspring genotype exerts an influence on the level of maternal investment and affects maternal fecundity through higher rates of embryo abortion and lower numbers of full-term offspring. Our results show that the prerequisites for parent–offspring conflict to be a potent evolutionary force in poeciliid fish are present in H. formosa. However, determining whether this conflict has shaped maternal investment in nature will require disentangling any effects of conflict from those of several ecological factors that are themselves correlated with the expected intensity of conflict.     

Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range

Understanding the mechanisms by which climate change will affect animal populations is vital for adaptive management. Many studies have described changes in the timing of biological events, which can produce phenological mismatch. Direct effects on prey abundance might also be important, but have rarely been studied. We examine the likely importance of variation in prey abundance in driving the demographics of a European golden plover ( Pluvialis apricaria ) population at its southern range margin. Previous studies have correlated plover productivity with the abundance of their adult cranefly (Tipulidae) prey, and modelled the phenology of both plover breeding and cranefly emergence in relation to temperature. Our analyses demonstrate that abundance of adult craneflies is correlated with August temperature in the previous year. Correspondingly, changes in the golden plover population are negatively correlated with August temperature 2 years earlier. Predictions of annual productivity, based on temperature-mediated reductions in prey abundance, closely match observed trends. Modelled variation in annual productivity for a future scenario of increasing August temperatures predicts a significant risk of extinction of the golden plover population over the next 100 years, depending upon the magnitude of warming. Direct effects of climate warming upon cranefly populations may therefore cause northward range contractions of golden plovers, as predicted by climate envelope modelling. Craneflies are an important food source for many northern and upland birds, and our results are likely to have wide relevance to these other species. Research into the potential for habitat management to improve the resilience of cranefly populations to high temperature should be an urgent priority.     

Differential reproductive responses to stress reveal the role of life-history strategies within a species

Life-history strategies describe that ‘slow’- in contrast to ‘fast’-living species allocate resources cautiously towards reproduction to enhance survival. Recent evidence suggests that variation in strategies exists not only among species but also among populations of the same species. Here, we examined the effect of experimentally induced stress on resource allocation of breeding seabirds in two populations with contrasting life-history strategies: slow-living Pacific and fast-living Atlantic black-legged kittiwakes. We tested the hypothesis that reproductive responses in kittiwakes under stress reflect their life-history strategies. We predicted that in response to stress, Pacific kittiwakes reduce investment in reproduction compared with Atlantic kittiwakes. We exposed chick-rearing kittiwakes to a short-term (3-day) period of increased exogenous corticosterone (CORT), a hormone that is released during food shortages. We examined changes in baseline CORT levels, parental care and effects on offspring. We found that kittiwakes from the two populations invested differently in offspring when facing stress. In response to elevated CORT, Pacific kittiwakes reduced nest attendance and deserted offspring more readily than Atlantic kittiwakes. We observed lower chick growth, a higher stress response in offspring and lower reproductive success in response to CORT implantation in Pacific kittiwakes, whereas the opposite occurred in the Atlantic. Our findings support the hypothesis that life-history strategies predict short-term responses of individuals to stress within a species. We conclude that behaviour and physiology under stress are consistent with trade-off priorities as predicted by life-history theory. We encourage future studies to consider the pivotal role of life-history strategies when interpreting inter-population differences of animal responses to stressful environmental events.     

Sub-saharan desertification and productivity are linked to hemispheric climate variability

Vegetation productivity and desertification in sub‐Saharan Africa may be influenced by global climate variability attributable to the North Atlantic Oscillation (NAO) and El Niño Southern Oscillation (ENSO). Combined and individual effects of the NAO and ENSO indices revealed that 75% of the interannual variation in the area of Sahara Desert was accounted for by the combined effects, with most variance attributable to the NAO. Effects were shown in the latitudinal variation on the 200 mm isocline, which was influenced mostly by the NAO. The combined indices explained much of the interannual variability in vegetation productivity in the Sahelian zone and southern Africa, implying that both the NAO and ENSO may be useful for monitoring effects of global climate change in sub‐Saharan Africa.     

Maternal grandmothers improve nutritional status and survival of children in rural Gambia

Hypotheses for the evolution of human female life-history characteristics have often focused on the social nature of human societies, which allows women to share the burden of childcare and provisioning amongst other members of their kin group. We test the hypothesis that child health and survival probabilities will be improved by the presence of kin using a longitudinal database from rural Gambia. We find that the only kin to improve the nutritional status of children significantly (apart from mothers) are maternal grandmothers, and that this is reflected in higher survival probabilities for children with living maternal grandmothers. There is also evidence that the reproductive status of the maternal grandmother influences child nutrition, with young children being taller in the presence of non-reproductive grandmothers than grandmothers who are still reproductively active. Paternal grandmothers and male kin, including fathers, have negligible impacts on the nutritional status and survival of children.     

The influence of maternal care in shaping human survival and fertility

The influence of maternal care on child survival has evolved throughout human history due to variation in altriciality, allocare, and maternal behaviors. Here, we study the impact of these factors on the force of selection acting on age-specific survival and fertility (measured with elasticity analysis) in a model that incorporates the dependence of child survival on maternal survival. Results reveal life-history changes that cannot be elucidated when considering child's survival independent of maternal survival: decrease of late fertility and increase of late survival, and concomitant decrease of early and late fertility. We also show that an increase of child altriciality in early humans might explain the main human life-history traits: a high life expectancy and postreproductive life; a long juvenile period and a higher, and narrowed, fertility at the peak of the reproductive period.     

Projected poleward shift of king penguins' (Aptenodytes patagonicus) foraging range at the Crozet Islands, southern Indian Ocean

Seabird populations of the Southern Ocean have been responding to climate change for the last three decades and demographic models suggest that projected warming will cause dramatic population changes over the next century. Shift in species distribution is likely to be one of the major possible adaptations to changing environmental conditions. Habitat models based on a unique long-term tracking dataset of king penguin (Aptenodytes patagonicus) breeding on the Crozet Islands (southern Indian Ocean) revealed that despite a significant influence of primary productivity and mesoscale activity, sea surface temperature consistently drove penguins' foraging distribution. According to climate models of the Intergovernmental Panel on Climate Change (IPCC), the projected warming of surface waters would lead to a gradual southward shift of the more profitable foraging zones, ranging from 25 km per decade for the B1 IPCC scenario to 40 km per decade for the A1B and A2 scenarios. As a consequence, distances travelled by incubating and brooding birds to reach optimal foraging zones associated with the polar front would double by 2100. Such a shift is far beyond the usual foraging range of king penguins breeding and would negatively affect the Crozet population on the long term, unless penguins develop alternative foraging strategies.