Insights from the first global population estimate of Weddell seals in Antarctica.

The Weddell seal is one of the best-studied marine mammals in the world, owing to a multidecadal demographic effort in the southernmost part of its range. Despite their occurrence around the Antarctic coastline, we know little about larger scale patterns in distribution, population size, or structure. We combined high-resolution satellite imagery from 2011, crowd-sourcing, and habitat modeling to report the first global population estimate for the species and environmental factors that influence its distribution. We estimated ~202,000 (95% confidence interval: 85,345 to 523,140) sub-adult and adult female seals, with proximate ocean depth and fast-ice variables as factors explaining spatial prevalence. Distances to penguin colonies were associated with seal presence, but only emperor penguin population size had a strong negative relationship. The small, estimated population size relative to previous estimates and the seals' nexus with trophic competitors indicates that a community ecology approach is required in efforts to monitor the Southern Ocean ecosystem.

Modeling changes in baleen whale seasonal abundance, timing of migration, and environmental variables to explain the sudden rise in entanglements in California.

We document changes in the number of sightings and timing of humpback (Megaptera novaeangliae), blue (Balaenoptera musculus), and gray (Eschrichtius robustus) whale migratory phases in the vicinity of the Farallon Islands, California. We hypothesized that changes in the timing of migration off central California were driven by local oceanography, regional upwelling, and basin-scale climate conditions. Using 24 years of daily whale counts collected from Southeast Farallon Island, we developed negative binomial regression models to evaluate trends in local whale sightings over time. We then used linear models to assess trends in the timing of migration, and to identify potential environmental drivers. These drivers included local, regional and basin-scale patterns; the latter included the El Niño Southern Oscillation, the Pacific Decadal Oscillation, and the North Pacific Gyre Oscillation, which influence, wind-driven upwelling, and overall productivity in the California Current System. We then created a forecast model to predict the timing of migration. Humpback whale sightings significantly increased over the study period, but blue and gray whale counts did not, though there was variability across the time series. Date of breeding migration (departure) for all species showed little to no change, whereas date of migration towards feeding areas (arrival) occurred earlier for humpback and blue whales. Timing was significantly influenced by a mix of local oceanography, regional, and basin-scale climate variables. Earlier arrival time without concomitant earlier departure time results in longer periods when blue and humpback whales are at risk of entanglement in the Gulf of the Farallones. We maintain that these changes have increased whale exposure to pot and trap fishery gear off the central California coast during the spring, elevating the risk of entanglements. Humpback entanglement rates were significantly associated with increased counts and early arrival in central California. Actions to decrease the temporal overlap between whales and pot/trap fishing gear, particularly when whales arrive earlier in warm water years, would likely decrease the risk of entanglements.

Modeling predator and prey hotspots: Management implications of baleen whale co-occurrence with krill in Central California.

As global ocean-bound commerce increases, managing human activities has become important in reducing conflict with threatened wildlife. This study investigates environmental factors determining abundance and distribution of blue whales (Balaenoptera musculus), humpback whales (Megaptera novaeangliae) and their prey (Euphausia pacifica and Thysanoessa spinifera) in central California. We provide insights into environmental drivers of the ecology and distribution of these species, model whale distributions and determine coincident hotspots of whales and their prey that will help decrease human threats to whales and protect critical feeding habitat. We developed separate predictive models of whale abundances (using negative binomial regression on count data) and krill abundance (using a two-part hurdlemodel combining logistic and negative binomial regressions) over a 14 year period (2004-2017). Variables included in situ surface and midwater oceanographic measures (temperature, salinity, and fluorescence), basin-scale climate indices, and bathymetric- and distance-related data. Predictions were applied to 1 km2 cells spanning the study area for May, June, July, and September during each of the 14 years of surveys to identify persistent distribution patterns. Both whales and krill were found to consistently use the northeast region of Cordell Bank, the Farallon Escarpment, and the shelf-break waters. The main identified blue whale hotspots were also krill hotspots, while co-occurrence was more limited and varied seasonally for humpback whales and krill. These results are valuable in identifying patterns in important areas of ecological interaction to assist management of whales. Areas north of Cordell Bank are of particular management concern since they overlap with the end of the San Francisco Bay northern shipping lane. Our findings can help decrease threats to whales, particularly in important foraging areas, by supporting implementation of vessel management and informing potential conflicts with other human uses.

Modeling Nonresident Seabird Foraging Distributions to Inform Ocean Zoning in Central California.

Seabird aggregations at sea have been shown to be associated with concentrations of prey. Previous research identified Central California as a highly used foraging area for seabirds, with locally breeding seabirds foraging close to their colonies on Southeast Farallon Island. Herein, we focus on nonresident (i.e. non-locally breeding) seabird species off of Central California. We hypothesized that high-use foraging areas for nonresident seabirds would be influenced by oceanographic and bathymetric factors and that spatial and temporal distributions would be similar within planktivorous and generalist foraging guilds but would differ between them. With data collected by the Applied California Current Ecosystem Studies (ACCESS) partnership during cruises between April and October from 2004-2013, we developed generalized linear models to identify high-use foraging areas for each of six nonresident seabird species. The four generalist species are Phoebastria nigripes (black-footed albatross), Ardenna griseus (sooty shearwater), Ardenna creatopus (pink-footed shearwater), and Fulmarus glacialis (northern fulmar). The two planktivorous species are Phalaropus lobatus (red-necked phalarope) and Phalaropus fulicarius (red phalarope). Sea surface temperature was significant for generalist species and sea surface salinity was important for planktivorous species. The distance to the 200-m isobath was significant in five of six models, Pacific Decadal Oscillation with a 3-month lag in four models, and sea surface fluorescence, the distance to Cordell Bank, and depth in three models. We did not find statistically significant differences between distributions of individual seabird species within a foraging guild or between guilds, with the exception of the sooty shearwater. Model results for a multi-use seabird foraging area highlighted the continental shelf break, particularly within the vicinity of Cordell Bank, as the highest use areas as did Marxan prioritization. Our research methods can be implemented elsewhere to identify critical habitat that needs protection as human development pressures continue to expand to the ocean.

Coping with the loss of large, energy-dense prey: a potential bottleneck for Weddell Seals in the Ross Sea.

Extraction of Antarctic toothfish (Dissostichus mawsoni) in the Ross Sea began in 1997, following a management plan that targets the largest fish with a goal of reducing the spawning biomass by 50% over 35 yr. We investigate the potential long-term consequences of the reduced availability of this prey for Weddell seals (Leptonychotes weddellii). Energy demands in seals are acute, especially immediately following lactation, when females must recover substantial mass and cope with molting costs. We tested the hypothesis that toothfish are critically important for adult female seals during this period. Toothfish body mass is three orders of magnitude greater, and its energy density nearly double that of the most common seal prey, Antarctic silverfish (Pleuragramma antarcticum). Reduction or elimination of toothfish consumption could impair a female's ability to sufficiently recover and successfully produce a pup in the following pupping season. Our goals are to (1) illustrate mechanisms and conditions whereby toothfish depletion might plausibly affect seal population trends; (2) identify measurable parameters of the seals' ecology that may help better understand the potential negative impact of toothfish depletion on seal populations; and (3) promote a precautionary management approach for the fishery that includes monitoring of seal populations We constructed a set of inter-linked models of seal diving behavior, physiological condition, and demography based on existing information. We evaluate the effect of the following factors on seal mass recovery and intrinsic population growth rates: fishery depletion rate, daily diving limits, probability of a successful dive, and body mass recovery target. We show that loss of toothfish has the greatest potential impact on seal populations' growth rate. Under some scenarios, populations may decrease at >10% per year. Critical parameters to better understand fishery impacts include prevalence and size of toothfish in the seals' diet; the relationship between diet and the rate of mass recovery; and female breeding propensity in relation to body condition at the end of the molting period. Our results lend support to concerns about the potential negative impact of toothfish extraction in the Ross Sea; and to advocate for a precautionary management approach by the fishery.

Spatial Distribution and Temporal Patterns of Cassin's Auklet Foraging and Their Euphausiid Prey in a Variable Ocean Environment.

Krill (Euphausiids) play a vital ecosystem role in many of the world's most productive marine regions, providing an important trophic linkage. We introduce a robust modeling approach to link Cassin's auklet (Ptychoramphus aleuticus) abundance and distribution to large-scale and local oceanic and atmospheric conditions and relate these patterns to similarly modeled distributions of an important prey resource, krill. We carried out at-sea strip transect bird surveys and hydroacoustic assessments of euphausiids (2004-2013). Data informed separate, spatially-explicit predictive models of Cassin's auklet abundance (zero-inflated negative binomial regression) and krill biomass (two-part model) based on these surveys. We established the type of prey responsible for acoustic backscatter by conducting net tows of the upper 50 m during surveys. We determined the types of prey fed to Cassin's auklet chicks by collecting diet samples from provisioning adults. Using time-depth-recorders, we found Cassin's auklets utilized consistent areas in the upper water column, less than 30 m, where krill could be found (99.5% of dives were less than 30 m). Birds primarily preyed upon two species of euphausiids, Euphausia pacifica and Thysanoessa spinifera, which were available in the upper water column. Cassin's auklet abundance was best predicted by both large scale and localized oceanic processes (upwelling) while krill biomass was best predicted by local factors (temperature, salinity, and fluorescence) and both large scale and localized oceanic processes (upwelling). Models predicted varying krill and bird distribution by month and year. Our work informs the use of Cassin's auklet as a valuable indicator or krill abundance and distribution and strengthens our understanding of the link between Cassin's auklet and its primary prey. We expect future increases in frequency and magnitude of anomalous ocean conditions will result in decreased availability of krill leading to declines in the Farallon Islands population of Cassin's auklets.

Using seabird habitat modeling to inform marine spatial planning in central California's National Marine Sanctuaries.

Understanding seabird habitat preferences is critical to future wildlife conservation and threat mitigation in California. The objective of this study was to investigate drivers of seabird habitat selection within the Gulf of the Farallones and Cordell Bank National Marine Sanctuaries to identify areas for targeted conservation planning. We used seabird abundance data collected by the Applied California Current Ecosystem Studies Program (ACCESS) from 2004-2011. We used zero-inflated negative binomial regression to model species abundance and distribution as a function of near surface ocean water properties, distances to geographic features and oceanographic climate indices to identify patterns in foraging habitat selection. We evaluated seasonal, inter-annual and species-specific variability of at-sea distributions for the five most abundant seabirds nesting on the Farallon Islands: western gull (Larus occidentalis), common murre (Uria aalge), Cassin's auklet (Ptychorampus aleuticus), rhinoceros auklet (Cerorhinca monocerata) and Brandt's cormorant (Phalacrocorax penicillatus). The waters in the vicinity of Cordell Bank and the continental shelf east of the Farallon Islands emerged as persistent and highly selected foraging areas across all species. Further, we conducted a spatial prioritization exercise to optimize seabird conservation areas with and without considering impacts of current human activities. We explored three conservation scenarios where 10, 30 and 50 percent of highly selected, species-specific foraging areas would be conserved. We compared and contrasted results in relation to existing marine protected areas (MPAs) and the future alternative energy footprint identified by the California Ocean Uses Atlas. Our results show that the majority of highly selected seabird habitat lies outside of state MPAs where threats from shipping, oil spills, and offshore energy development remain. This analysis accentuates the need for innovative marine spatial planning efforts and provides a foundation on which to build more comprehensive zoning and management in California's National Marine Sanctuaries.

Where the wild things are: predicting hotspots of seabird aggregations in the California Current System.

Marine protected areas (MPAs) provide an important tool for conservation of marine ecosystems. To be most effective, these areas should be strategically located in a manner that supports ecosystem function. To inform marine spatial planning and support strategic establishment of MPAs within the California Current System, we identified areas predicted to support multispecies aggregations of seabirds ("hotspots"). We developed habitat-association models for 16 species using information from at-sea observations collected over an 11-year period (1997-2008), bathymetric data, and remotely sensed oceanographic data for an area from north of Vancouver Island, Canada, to the USA/Mexico border and seaward 600 km from the coast. This approach enabled us to predict distribution and abundance of seabirds even in areas of few or no surveys. We developed single-species predictive models using a machine-learning algorithm: bagged decision trees. Single-species predictions were then combined to identify potential hotspots of seabird aggregation, using three criteria: (1) overall abundance among species, (2) importance of specific areas ("core areas") to individual species, and (3) predicted persistence of hotspots across years. Model predictions were applied to the entire California Current for four seasons (represented by February, May, July, and October) in each of 11 years. Overall, bathymetric variables were often important predictive variables, whereas oceanographic variables derived from remotely sensed data were generally less important. Predicted hotspots often aligned with currently protected areas (e.g., National Marine Sanctuaries), but we also identified potential hotspots in Northern California/Southern Oregon (from Cape Mendocino to Heceta Bank), Southern California (adjacent to the Channel Islands), and adjacent to Vancouver Island, British Columbia, that are not currently included in protected areas. Prioritization and identification of multispecies hotspots will depend on which group of species is of highest management priority. Modeling hotspots at a broad spatial scale can contribute to MPA site selection, particularly if complemented by fine-scale information for focal areas.

Climate and demography of the planktivorous Cassin's auklet Ptychoramphus aleuticus off northern California: implications for population change.

1. We performed demographic analyses on Cassin's auklet Ptychoramphus aleuticus, a zooplanktivorous seabird inhabiting the variable California Current System, to understand how temporal environmental variability influences population dynamics. 2. We used capture-recapture data from 1986 to 2002 to rank models of interannual variation in survival, breeding propensity, breeding success, and recruitment. 3. All demographic parameters exhibited temporal variability. Interannual variation in survival was best modelled as a nonlinear function of the winter Southern Oscillation Index (SOI). Breeding propensity was best modelled as a threshold function of local sea surface temperature. Breeding success and recruitment were best modelled with year-dependent annual variation. 4. Changes in the SOI force El Niño/La Niña events, which in turn alter prey availability to seabirds in this system. Demographic responses varied during El Niños/La Niñas. Survival diminished substantially during the 1997-98 El Niño event, while breeding propensity was affected during both the 1992 and 1998 El Niños. Breeding success was reduced during the 1992, 1993, and 1998 El Niños, but was unusually high in 2002. Recruitment was higher at the beginning and end of this time-series. 5. While demographic responses varied interannually, parameter values covaried in a positive fashion, a situation conducive to rapid population change. During the 11 years study period, the Farallon auklet breeding population declined at 6.05 +/- 0.80% (SE) per year, a cumulative decline of 49.7%. This study demonstrates how climate variability has influenced key demographic processes for this diminished marine bird population.

Altitudinal variation in parental energy expenditure by white-crowned sparrows.

We used the doubly labeled water technique to measure daily energy expenditure (DEE) during the incubation and feeding nestling stages in two populations of white-crowned sparrows (Zonotrichia leucophrys) - one montane and migratory, the other coastal and sedentary - that differ in thermal environment and clutch size. We assessed the birds' thermal environment by continuously monitoring (among other variables) operative temperature and wind speed both in the open and within bushes and willow thickets occupied by sparrows. From these measurements, we derived several estimates of the birds' thermal environment, including standard operative temperature (T(es)). Shade air temperature and T(es) averaged 6.6 and 10.3 degrees C lower, respectively, at the montane study site during DEE measurements. The montane population's DEE averaged 24% higher than that of the sea-level population (103.6+/-12.2 versus 83.7+/-9.6 kJ day(-1); means +/- S.D., N=31 and 22, respectively), reflecting both its larger brood size (3.7 versus 2.9) and the colder environment. The DEE:BMR ratio was lowest in the sea-level population (2.1 versus 2.6), but neither population worked to their physiological capacity to produce young. DEE was significantly correlated with temperature across populations, with T(es) explaining 42% of the variation in DEE. Statistically removing the effect of temperature by adjusting DEE to a common temperature reduced the difference in DEE between populations by 34% to 87.7 and 100.8 kJ day(-1), respectively, for sea-level and montane populations. Basal and resting metabolic rates were similar in both populations, implying that greater activity in the montane population accounted for its higher temperature-adjusted DEE. Our results indicate that the thermal context within which behavior occurs can significantly affect interindividual variation in DEE. Attempts to assess reproductive effort by measuring DEE should therefore account explicitly for the effect of temperature.

THE CONSEQUENCES OF BROOD SIZE FOR BREEDING BLUE TITS. III. MEASURING THE COST OF REPRODUCTION: SURVIVAL, FUTURE FECUNDITY, AND DIFFERENTIAL DISPERSAL.

To determine how the cost of reproduction varies with brood size, a population of blue tits (Parus caeruleus) breeding in Wytham Wood, England, was manipulated over a three year period. Two hundred sixteen pairs were randomly assigned 3, 6, 9, 12, or 15 nestlings; nestlings were exchanged soon after hatching. Survival of adult females (as measured by the proportion recaptured in the following winter and/or spring) declined significantly with increasing brood size in two out of three years; there was significant year-to-year variation in the relationship of recapture rate to brood size. Mean female recapture rates (averaged over the three years) declined in a linear fashion (P < 0.01). There was no significant linear or curvilinear relationship between male-recapture rate and brood size in any of the three years nor was there a significant linear or curvilinear relationship for the data averaged over the three years. Nevertheless, recapture proportions for males differed significantly with respect to brood size (χ2 test, P < 0.05). The possibility that experimental brood size influences subsequent dispersal (and therefore biases measures of survival based on recapture rates to differing degrees) was examined by comparing distances moved by breeding adults from one year to the next. There was no relationship between brood size and dispersal distance within the study area for either sex, except that females given broods of three were significantly more likely to move more than 300 m than were those given broods of 6-15 young. Both males and females showed evidence of a cost with respect to future fecundity: as brood size increased, the number of surviving offspring produced in the following year decreased from 1.5-1.6 (for adults that had reared 3-6 young) to 0.4 (for those that had reared 15 young). The relationship of future reproductive success to experimental brood size did not differ among years or between the sexes. The number of eggs laid and number of young hatched in year n + 1 did not differ significantly with respect to brood size in year n; rather, differences in future fecundity reflected differences in the survival prospects of young reared in year n + 1.

Feeding frequencies of nestling blue tits (Parus caeruleus): costs, benefits and a model of optimal feeding frequency.

1. To examine the ultimate factors determining the frequency with which male and female blue tits (Parus caeruleus) feed their nestlings, I manipulated the brood sizes of 34 pairs breeding in Wytham Woods, England, and recorded automatically the number of visits when the young were 11-13 days of age. At this time adults were caught and weighed and the nestlings weighed. 2. The number of visits/brood increased linearly with brood size, with no evidence of an upper limit to brood feeding frequency. The number of visits·nestlings-1·day-1 decreased from 138 to 108 as brood size increased from 3 to 6, but levelled off at 68-87 visits/nestling as brood size increased from 8 to 15. 3. Female weight-loss during the nestling period increased linearly as the feeding frequency of the brood increased: those females were lightest that fed their young most often; this is of some significance since I previously showed that the lightest females are least likely to survive to the following year. The correlations of feeding frequency with brood size and feeding frequency with female weight are sufficient to account for the commonly observed correlation of brood size and female weight. No relationship between feeding frequency and male weight dynamics was apparent. Feeding frequency of the brood increased as mean nestling weight decreased. Apparently, parents compensate, in part, for low nestling weight by increasing feeding frequency. Low nestling weight, in turn, has proven detrimental to post-fledging survival. 4. Lack's (1947, 1954) hypothesis, that the decline in visits/nestling as brood size increases reflects the inability of parents to sustain high brood feeding rates, is refuted by the observations made in this and a large number of other studies. Royama's (1966) hypothesis that the observed decline in visits/nestling reflects a decrease in energy needed to maintain homeothermy, receives some support. Behavioral and physiological studies, however, suggest that the "Royama effect" is limited to small broods (c. four or fewer young). 5. I propose, instead, a model of optimal feeding frequency, which accounts for observations made by myself and others, and which incorporates Royama's hypothesis as well. The model assumes that parents optimize their investment in their young. On the one hand, an increase in feeding frequency means more food for the young, which enhances offspring survival ("benefit"); on the other hand, increased feeding frequency is "costly" to the parents (reduces parental survivorship). Natural selection favors breeding individuals that maximize the difference between benefits (to the offpsring) and costs (to the parents). The model accounts for many aspects of blue tit reproductive biology, including the observed relationship between nestling weight and brood size and the observation that parent reproductive costs increase with brood size.