Russell W. Bradley

Timing is everything: flexible phenology and shifting selection in a colonial seabird

1. In order to reproduce successfully in a temporally varying environment, iteroparous animals must exhibit considerable behavioural flexibility across their lifetimes. By adjusting timing of breeding each year, parents can ensure optimal overlap between the energy intensive period of offspring production and the seasonal peak in favourable environmental conditions, thereby increasing their chances of successfully rearing young. 2. Few studies investigate variation among individuals in how they respond to fluctuating conditions, or how selection acts on these individual differences, but this information is essential for understanding how populations will cope with rapid environmental change. 3. We explored inter-annual trends in breeding time and individual responses to environmental variability in common guillemots Uria aalge, an important marine top predator in the highly variable California Current System. Complex, nonlinear relationships between phenology and oceanic and climate variables were found at the population level. Using a novel application of a statistical technique called random regression, we showed that individual females responded in a nonlinear fashion to environmental variability, and that reaction norm shape differed among females. 4. The pattern and strength of selection varied substantially over a 34-year period, but in general, earlier laying was favoured. Females deviating significantly from the population mean laying date each year also suffered reduced breeding success, with the strength of nonlinear selection varying in relation to environmental conditions. 5. We discuss our results in the wider context of an emerging literature on the evolutionary ecology of individual-level plasticity in the wild. Better understanding of how species-specific factors and local habitat features affect the timing and success of breeding will improve our ability to predict how populations will respond to climate change.

As the egg turns: monitoring egg attendance behavior in wild birds using novel data logging technology.

Egg turning is unique to birds and critical for embryonic development in most avian species. Technology that can measure changes in egg orientation and temperature at fine temporal scales (1 Hz) was neither readily available nor small enough to fit into artificial eggs until recently. Here we show the utility of novel miniature data loggers equipped with 3-axis (i.e., triaxial) accelerometers, magnetometers, and a temperature thermistor to study egg turning behavior in free-ranging birds. Artificial eggs containing egg loggers were deployed in the nests of three seabird species for 1–7 days of continuous monitoring. These species (1) turned their eggs more frequently (up to 6.5 turns h−1) than previously reported for other species, but angular changes were often small (1–10° most common), (2) displayed similar mean turning rates (ca. 2 turns h−1) despite major differences in reproductive ecology, and (3) demonstrated distinct diurnal cycling in egg temperatures that varied between 1.4 and 2.4°C. These novel egg loggers revealed high-resolution, three-dimensional egg turning behavior heretofore never measured in wild birds. This new form of biotechnology has broad applicability for addressing fundamental questions in avian breeding ecology, life history, and development, and can be used as a tool to monitor birds that are sensitive to disturbance while breeding.

Recruitment of Cassin's Auklet (Ptychoramphus aleuticus): Individual Age and Parental Age Effects

ABSTRACT. Recruitment is an important aspect of life history that is difficult to estimate without long-term data. We used 28 years of data to create multistate capture—mark—recapture models with maturity status as state to provide estimates of age-dependent probabilistic recruitment (transition from juvenile to breeder state) of Cassins Auklets (Ptychoramphus aleuticus) breeding at Southeast Farallon Island, California. We found strong evidence for age effects in recruitment and survival, but not in breeding propensity. We also examined parental-age effects on return rate of offspring. Although an average of 70% of a cohort begins breeding by age 4, fledglings from 2- to 4-year-old mothers were less likely to become breeders than fledglings produced by older mothers. This may be attributable to age-related differences in egg production and maternal offspring-rearing abilities that could contribute to differences in fledgling recruitment rates. Eggs of younger mothers had lower volume and their fledglings had lower mass than those of older mothers. We suggest that researchers, managers, and population modelers consider the possibility that young breeding birds that successfully fledge young do not always produce viable offspring that will become breeders.

Dynamics of White Shark Prédation on Pinnipeds in California: Effects of Prey Abundance

Abstract To assess long-term temporal trends in White Shark (Carcharadon carcharias) predation, and examine the relationship between C. carcharias and pinnipeds, we examined a 17-year (1988–2004) data set of daily shark attack surveys from Southeast Farallon Island (SEFI), California. We modeled within-season and among-year variation in the number of observed shark attacks at SEFI. Within-season, daily probability of attack was affected by hours of effort, date, tide height, and pinniped abundance. The seasonal peak in shark predation did not vary inter-annually. Daily probability of attack was positively correlated with pinniped abundance and negatively correlated with tide height. After controlling for within-season effects, 51% of annual variation in the number of observed shark attacks was explained by an inverse function of the abundance of Northern Elephant Seals (Mirounga angustirostris). The addition of shark abundance to the model did not significantly increase the models descriptive power. This suggests a functional response where predation by C. carcharias increased with abundance of M. angustirostris, but plateaued once a critical density of M. angustirostris were available, either due to limitations of prey handling or satiation. We detected no temporal trend in annual number of observed shark attacks. Therefore, C. carcharias depends on M. angustirostris as a critical food source and will increase or decrease predation rates depending upon current seal populations.

Life span and reproductive cost explain interspecific variation in the optimal onset of reproduction

Fitness can be profoundly influenced by the age at first reproduction (AFR), but to date the AFR–fitness relationship only has been investigated intraspecifically. Here, we investigated the relationship between AFR and average lifetime reproductive success (LRS) across 34 bird species. We assessed differences in the deviation of the Optimal AFR (i.e., the species‐specific AFR associated with the highest LRS) from the age at sexual maturity, considering potential effects of life history as well as social and ecological factors. Most individuals adopted the species‐specific Optimal AFR and both the mean and Optimal AFR of species correlated positively with life span. Interspecific deviations of the Optimal AFR were associated with indices reflecting a change in LRS or survival as a function of AFR: a delayed AFR was beneficial in species where early AFR was associated with a decrease in subsequent survival or reproductive output. Overall, our results suggest that a delayed onset of reproduction beyond maturity is an optimal strategy explained by a long life span and costs of early reproduction. By providing the first empirical confirmations of key predictions of life‐history theory across species, this study contributes to a better understanding of life‐history evolution.

Growth, Age at Maturity, and Age-Specific Survival of the Arboreal Salamander (Aneides lugubris) on Southeast Farallon Island, California

Abstract Growth, age at maturity, and survival are life-history parameters that provide important information for understanding population dynamics. We modeled growth and age at maturity for an island population of Arboreal Salamanders, Aneides lugubris, using snout–vent length (SVL) growth intervals from a 4-yr capture–mark–recapture study fit to the von Bertalanffy growth interval model. We estimated annual survival as a function of SVL using a multistate open robust design model, and computed age-specific survival using results from the von Bertalanffy growth model. Arboreal Salamanders have indeterminate growth that slows with age from hatchling size (24.4-mm SVL) to the mean adult (asymptotic) size of 66.0-mm SVL. Age at maturity is 2.69 yr, and average adult age is 8–11 yr. Annual survival increased with age from 0.363 in age 0 to 0.783 in ages >4 yr. Our results provide the first estimates of life-history parameters for this species and indicate similarities to other terrestrial salamanders from low-elevation Mediterranean climates.

Shifting Effects of Ocean Conditions on Survival and Breeding Probability of a Long-Lived Seabird

With a rapidly changing climate, there is an increasing need to predict how species will respond to changes in the physical environment. One approach is to use historic data to estimate the past influence of environmental variation on important demographic parameters and then use these relationships to project the abundance of a population or species under future climate scenarios. However, as novel climate conditions emerge, novel species responses may also appear. In some systems, environmental conditions beyond the range of those observed during the course of most long-term ecological studies are already evident. Yet little attention has been given to how these novel conditions may be influencing previously established environment–species relationships. Here, we model the relationships between ocean conditions and the demography of a long-lived seabird, Brandt’s cormorant (Phalacrocorax penicillatusI), in central California and show that these relationships have changed in recent years. Beginning in 2007/2008, the response of Brandt’s cormorant, an upper trophic level predator, to ocean conditions shifted, resulting in lower than predicted survival and breeding probability. Survival was generally less variable than breeding probability and was initially best predicted by the basin-scale forcing of the El Niño Southern Oscillation rather than local ocean conditions. The shifting response of Brandt’s cormorant to ocean conditions may be just a proximate indication of altered dynamics in the food web and that important forage fish are not responding to the physical ocean environment as expected. These changing relationships have important implications for our ability to project the effects of future climate change for species and communities.

Global phenological insensitivity to shifting ocean temperatures among seabirds

Reproductive timing in many taxa plays a key role in determining breeding productivity1, and is often sensitive to climatic conditions2. Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey3. This is of particular concern for higher-trophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers4. However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction5. Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1) or in response to sea surface temperature (SST) (−0.272 days °C−1) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources2.Time of reproduction may be altered as the climate changes. For seabirds, it is shown that there has not been an adjustment in timing as the climate changes and the sea surface warms. This lack of plasticity could result in a mismatch with food resources.

Changing environmental spectra influence age-structured populations: increasing ENSO frequency could diminish variance and extinction risk in long-lived seabirds

As global climate changes, there is increasing need to understand how changes in the frequencies of environmental variability affect populations. Age-structured populations have recently been shown to filter specific frequencies of environmental variability, favoring generational frequencies, and very low frequencies, a phenomenon known as cohort resonance. However, there has been little exploration of how changes in the spectra of environmental signals will affect the stability and persistence of age-structured populations. To examine this issue, we analyzed a likely example to show how changes in the frequency of an influential climate phenomenon, the El Niño-Southern Oscillation (ENSO), could affect a marine bird population. We used a density-dependent, age-structured population model to calculate the transfer function (i.e., the frequency-dependent sensitivity) of Brandt’s cormorant (Phalacrocorax penicillatus), a representative marine bird species known to be influenced by ENSO. We then assessed how the population would be affected by ENSO forcing that was doubled and halved in frequency. The transfer function indicated this population is most sensitive to variance at low frequencies, but does not exhibit the sensitivity to generational frequencies (cohort resonance) observed in shorter-lived species. Doubling the frequency of ENSO unexpectedly resulted in higher mean adult population abundance, lower variance, and lower probability of extinction, compared to forcing with the historical or reduced ENSO frequency. Our results illustrate how long-lived species with environmentally driven variability in recruitment, including many species of marine birds and fish, may respond in counterintuitive ways to anticipated changes in environmental variability.

Increased reproductive investment associated with greater survival and longevity in Cassin's auklets

Individuals increase lifetime reproductive output through a trade-off between investment in future survival and immediate reproductive success. This pattern may be obscured in certain higher quality individuals that possess greater reproductive potential. The Cassins auklet (Ptychoramphus aleuticus) is a long-lived species where some individuals exhibit greater reproductive ability through a behaviour called double brooding. Here, we analyse 32 years of breeding histories from marked known-age auklets to test whether double brooding increases lifetime fitness despite the increased mortality and reduced lifespan higher reproductive effort would be expected to incur. Multistate mark–recapture modelling revealed that double brooding was strongly positively associated with higher annual survival and longevity. The mean (95% confidence interval) apparent survival was 0.69 (0.21, 0.91) for individuals that executed a single brood and 0.96 (0.84, 0.99) for those that double-brooded. Generalized linear mixed models indicated individuals that attempted multiple double broods over their lifetime were able to produce on average seven times as many chicks and live nearly 6 years longer than birds that never attempted a double brood. We found that high-quality individuals exhibited both increased reproductive effort and longevity, where heterogeneity in individual quality masked expected life-history trade-offs.