Going with the floe: Sea-ice movement affects distance and destination during Adélie penguin winter movements.

Seasonal migration, driven by shifts in annual climate cycles and resources, is a key part of the life history and ecology of species across taxonomic groups. By influencing the amount of energy needed to move, external forces such as wind and ocean currents are often key drivers of migratory pathways exposing individuals to varying resources, environmental conditions, and competition pressures impacting individual fitness and population dynamics. Although wildlife movements in connection with wind and ocean currents are relatively well understood, movements within sea-ice fields have been much less studied, despite sea ice being an integral part of polar ecology. Adélie penguins (Pygoscelis adeliae) in the southern Ross Sea, Antarctica, currently exist at the southernmost edge of their range and undergo the longest (~12,000 km) winter migration known for the species. Within and north of the Ross Sea, the Ross Gyre drives ocean circulation and the large-scale movement of sea ice. We used remotely sensed sea-ice movement data together with geolocation-based penguin movement data to test the hypothesis that penguins use gyre-driven sea-ice movement to aid their migration. We found that penguins traveled greater distances when their movement vectors were aligned with those of sea ice (i.e., ice support) and the amount of ice support received depended on which route a penguin took. We also found that birds that took an eastern route traveled significantly further north in two of the 3 years we examined, coinciding with higher velocities of sea ice in those years. We compare our findings to patterns observed in migrating species that utilize air or water currents for their travel and with other studies showing the importance of ocean/sea-ice circulation patterns to wildlife movement and life history patterns within the Ross Sea. Changes in sea ice may have consequences not only for energy expenditure but, by altering migration and movement pathways, to the ecological interactions that exist in this region.

Sea ice concentration decline in an important Adélie penguin molt area.

Unlike in many polar regions, the spatial extent and duration of the sea ice season have increased in the Ross Sea sector of the Southern Ocean during the satellite era. Simultaneously, populations of Adélie penguins, a sea ice obligate, have been stable or increasing in the region. Relationships between Adélie penguin population growth and sea ice concentration (SIC) are complex, with sea ice driving different, sometimes contrasting, demographic patterns. Adélie penguins undergo a complete molt annually, replacing all their feathers while fasting shortly after the breeding season. Unlike most penguin species, a majority of Adélies are thought to molt on sea ice, away from the breeding colonies, which makes this period particularly difficult to study. Here, we evaluate the hypothesis that persistent areas of high SIC provide an important molting habitat for Adélie penguins. We analyzed data from geolocating dive recorders deployed year-round on 195 adult penguins at two colonies in the Ross Sea from 2017 to 2019. We identified molt by detecting extended gaps in postbreeding diving activity and used associated locations to define two key molting areas. Remotely sensed data indicated that SIC during molt was anomalously low during the study and has declined in the primary molt area since 1980. Further, annual return rates of penguins to breeding colonies were positively correlated with SIC in the molt areas over 20 y. Together these results suggest that sea ice conditions during Adélie penguin molt may represent a previously underappreciated annual bottleneck for adult survival.

High-resolution recording of foraging behaviour over multiple annual cycles shows decline in old Adélie penguins' performance.

Age-related variation in foraging performance can result from both within-individual change and selection processes. These mechanisms can only be disentangled by using logistically challenging long-term, longitudinal studies. Coupling a long-term demographic data set with high-temporal-resolution tracking of 18 Adélie penguins (Pygoscelis adeliae, age 4-15 yrs old) over three consecutive annual cycles, we examined how foraging behaviour changed within individuals of different age classes. Evidence indicated within-individual improvement in young and middle-age classes, but a significant decrease in foraging dive frequency within old individuals, associated with a decrease in the dive descent rate. Decreases in foraging performance occurred at a later age (from 12-15 yrs old to 15-18 yrs old) than the onset of senescence predicted for this species (9-11 yrs old). Foraging dive frequency was most affected by the interaction between breeding status and annual life-cycle periods, with frequency being highest during returning migration and breeding season and was highest overall for successful breeders during the chick-rearing period. Females performed more foraging dives per hour than males. This longitudinal, full annual cycle study allowed us to shed light on the changes in foraging performance occurring among individuals of different age classes and highlighted the complex interactions among drivers of individual foraging behaviour.

The influence of subcolony-scale nesting habitat on the reproductive success of Adélie penguins.

Group-size variation is common in colonially breeding species, including seabirds, whose breeding colonies can vary in size by several orders of magnitude. Seabirds are some of the most threatened marine taxa and understanding the drivers of colony size variation is more important than ever. Reproductive success is an important demographic parameter that can impact colony size, and it varies in association with a number of factors, including nesting habitat quality. Within colonies, seabirds often aggregate into distinct groups or subcolonies that may vary in quality. We used data from two colonies of Adélie penguins 73 km apart on Ross Island, Antarctica, one large and one small to investigate (1) How subcolony habitat characteristics influence reproductive success and (2) How these relationships differ at a small (Cape Royds) and large (Cape Crozier) colony with different terrain characteristics. Subcolonies were characterized using terrain attributes (elevation, slope aspect, slope steepness, wind shelter, flow accumulation), as well group characteristics (area/size, perimeter-to-area ratio, and proximity to nest predators). Reproductive success was higher and less variable at the larger colony while subcolony characteristics explained more of the variance in reproductive success at the small colony. The most important variable influencing subcolony quality at both colonies was perimeter-to-area ratio, likely reflecting the importance of nest predation by south polar skuas along subcolony edges. The small colony contained a higher proportion of edge nests thus higher potential impact from skua nest predation. Stochastic environmental events may facilitate smaller colonies becoming "trapped" by nest predation: a rapid decline in the number of breeding individuals may increase the proportion of edge nests, leading to higher relative nest predation and hindering population recovery. Several terrain covariates were retained in the final models but which variables, the shapes of the relationships, and importance varied between colonies.

Identification of a Novel Adélie Penguin Circovirus at Cape Crozier (Ross Island, Antarctica).

Understanding the causes of disease in Antarctic wildlife is crucial, as many of these species are already threatened by environmental changes brought about by climate change. In recent years, Antarctic penguins have been showing signs of an unknown pathology: a feather disorder characterised by missing feathers, resulting in exposed skin. During the 2018-2019 austral summer breeding season at Cape Crozier colony on Ross Island, Antarctica, we observed for the first time an Adélie penguin chick missing down over most of its body. A guano sample was collected from the nest of the featherless chick, and using high-throughput sequencing, we identified a novel circovirus. Using abutting primers, we amplified the full genome, which we cloned and Sanger-sequenced to determine the complete genome of the circovirus. The Adélie penguin guano-associated circovirus genome shares <67% genome-wide nucleotide identity with other circoviruses, representing a new species of circovirus; therefore, we named it penguin circovirus (PenCV). Using the same primer pair, we screened 25 previously collected cloacal swabs taken at Cape Crozier from known-age adult Adélie penguins during the 2014-2015 season, displaying no clinical signs of feather-loss disorder. Three of the 25 samples (12%) were positive for a PenCV, whose genome shared >99% pairwise identity with the one identified in 2018-2019. This is the first report of a circovirus associated with a penguin species. This circovirus could be an etiological agent of the feather-loss disorder in Antarctic penguins.

Impact of extreme drought and incentive programs on flooded agriculture and wetlands in California's Central Valley.

BACKGROUND: Between 2013 and 2015, a large part of the western United States, including the Central Valley of California, sustained an extreme drought. The Central Valley is recognized as a region of hemispheric importance for waterbirds, which use flooded agriculture and wetlands as habitat. Thus, the impact of drought on the distribution of surface water needed to be assessed to understand the effects on waterbird habitat availability. METHODS: We used remote sensing data to quantify the impact of the recent extreme drought on the timing and extent of waterbird habitat during the non-breeding season (July-May) by examining open water in agriculture (rice, corn, and other crops) and managed wetlands across the Central Valley. We assessed the influence of habitat incentive programs, particularly The Nature Conservancy's BirdReturns and The Natural Resources Conservation Service's Waterbird Habitat Enhancement Program (WHEP), at offsetting habitat loss related to drought. RESULTS: Overall, we found statistically significant declines in open water in post-harvest agriculture (45-80% declines) and in managed wetlands (39-60% declines) during the 2013-2015 drought compared to non-drought years during the period of 2000-2011. Crops associated with the San Joaquin Basin, specifically corn, as well as wetlands in that part of the Central Valley exhibited larger reductions in open water than rice and wetlands in the Sacramento Valley. Semi-permanent wetlands on protected lands had significantly lower (39-49%) open water in the drought years than those on non-protected lands while seasonal wetlands on protected lands had higher amounts of open water. A large fraction of the daily open water in rice during certain times of the year, particularly in the fall for BirdReturns (61%) and the winter for WHEP (100%), may have been provided through incentive programs which underscores the contribution of these programs. However, further assessment is needed to know how much the incentive programs directly offset the impact of drought in post-harvest rice by influencing water management or simply supplemented funding for activities that might have been done regardless. DISCUSSION: Our landscape analysis documents the significant impacts of the recent extreme drought on freshwater wetland habitats in the Central Valley, the benefits of incentive programs, and the value of using satellite data to track surface water and waterbird habitats. More research is needed to understand subsequent impacts on the freshwater dependent species that rely on these systems and how incentive programs can most strategically support vulnerable species during future extreme drought.

Unique genome organization of non-mammalian papillomaviruses provides insights into the evolution of viral early proteins.

The family Papillomaviridae contains more than 320 papillomavirus types, with most having been identified as infecting skin and mucosal epithelium in mammalian hosts. To date, only nine non-mammalian papillomaviruses have been described from birds (n = 5), a fish (n = 1), a snake (n = 1), and turtles (n = 2). The identification of papillomaviruses in sauropsids and a sparid fish suggests that early ancestors of papillomaviruses were already infecting the earliest Euteleostomi. The Euteleostomi clade includes more than 90 per cent of the living vertebrate species, and progeny virus could have been passed on to all members of this clade, inhabiting virtually every habitat on the planet. As part of this study, we isolated a novel papillomavirus from a 16-year-old female Adélie penguin (Pygoscelis adeliae) from Cape Crozier, Ross Island (Antarctica). The new papillomavirus shares ∼64 per cent genome-wide identity to a previously described Adélie penguin papillomavirus. Phylogenetic analyses show that the non-mammalian viruses (expect the python, Morelia spilota, associated papillomavirus) cluster near the base of the papillomavirus evolutionary tree. A papillomavirus isolated from an avian host (Northern fulmar; Fulmarus glacialis), like the two turtle papillomaviruses, lacks a putative E9 protein that is found in all other avian papillomaviruses. Furthermore, the Northern fulmar papillomavirus has an E7 more similar to the mammalian viruses than the other avian papillomaviruses. Typical E6 proteins of mammalian papillomaviruses have two Zinc finger motifs, whereas the sauropsid papillomaviruses only have one such motif. Furthermore, this motif is absent in the fish papillomavirus. Thus, it is highly likely that the most recent common ancestor of the mammalian and sauropsid papillomaviruses had a single motif E6. It appears that a motif duplication resulted in mammalian papillomaviruses having a double Zinc finger motif in E6. We estimated the divergence time between Northern fulmar-associated papillomavirus and the other Sauropsid papillomaviruses be to around 250 million years ago, during the Paleozoic-Mesozoic transition and our analysis dates the root of the papillomavirus tree between 400 and 600 million years ago. Our analysis shows evidence for niche adaptation and that these non-mammalian viruses have highly divergent E6 and E7 proteins, providing insights into the evolution of the early viral (onco-)proteins.

Improving effectiveness of systematic conservation planning with density data.

Systematic conservation planning aims to design networks of protected areas that meet conservation goals across large landscapes. The optimal design of these conservation networks is most frequently based on the modeled habitat suitability or probability of occurrence of species, despite evidence that model predictions may not be highly correlated with species density. We hypothesized that conservation networks designed using species density distributions more efficiently conserve populations of all species considered than networks designed using probability of occurrence models. To test this hypothesis, we used the Zonation conservation prioritization algorithm to evaluate conservation network designs based on probability of occurrence versus density models for 26 land bird species in the U.S. Pacific Northwest. We assessed the efficacy of each conservation network based on predicted species densities and predicted species diversity. High-density model Zonation rankings protected more individuals per species when networks protected the highest priority 10-40% of the landscape. Compared with density-based models, the occurrence-based models protected more individuals in the lowest 50% priority areas of the landscape. The 2 approaches conserved species diversity in similar ways: predicted diversity was higher in higher priority locations in both conservation networks. We conclude that both density and probability of occurrence models can be useful for setting conservation priorities but that density-based models are best suited for identifying the highest priority areas. Developing methods to aggregate species count data from unrelated monitoring efforts and making these data widely available through ecoinformatics portals such as the Avian Knowledge Network will enable species count data to be more widely incorporated into systematic conservation planning efforts.

Mejoría de la Efectividad de la Planeación Sistemática de la Conservación con Datos de Densidad Resumen La planeación sistemática de la conservación tiene como meta diseñar redes de áreas protegidas que cumplan con objetivos de conservación a lo largo de grandes paisajes. El diseño óptimo de estas redes de conservación se basa con mayor frecuencia en modelos de idoneidad de hábitat o probabilidad de occurrencia de especies, a pesar de la evidencia existente de que las predicciones de esos modelos pueden no estar fuertemente correlacionadas con la densidad de especies. Hipotetizamos que las redes de conservación diseñadas con las distribuciones de la densidad de especies conservan con mayor eficiencia a las poblaciones de todas las especies consideradas que las redes diseñadas con modelos de probabilidad de occurencia. Para probar esta hipótesis usamos el algoritmo Zonation de planeación de la conservación para evaluar los diseños de redes de conservación basados en la probabilidad de ocurrencia versus los modelos de densidad para 26 especies de aves terrestres en el noroeste del Pacífico en los Estados Unidos. Evaluamos la efectividad de cada red de conservación con base en las densidades pronosticadas de cada especie y la diversidad de especies pronosticada. Las clasificaciones de Zonation de los modelos de alta densidad protegieron a más individuos por especie cuando las redes protegieron el 10-40% de la más alta prioridad del paisaje. Comparado con los modelos basados en la densidad, los modelos basados en la ocurrencia protegieron a más individuos en el 50% más bajo de las áreas prioritarias de los paisajes. Las dos estrategias conservaron la diversidad de especies de formas similares: la diversidad pronosticada fue más alta en las localidades de alta prioridad en ambas redes de conservación. Concluimos que tanto los modelos de densidad como los de probabilidad de ocurrencia pueden ser útiles para establecer prioridades de conservación, pero que los modelos basados en la densidad son más adecuados para identificar las áreas de más alta prioridad. Desarrollar métodos para agregar datos de conteos de especies a partir de esfuerzos de monitoreo no relacionados y hacer que estos datos estén disponibles en portales eco-informáticos como la Avian Knowledge Network permitirá que los datos de conteos de especies se incorporen más ampliamente en esfuerzos de planeación sistemática de la conservación.

Evaluating tidal marsh sustainability in the face of sea-level rise: a hybrid modeling approach applied to San Francisco Bay.

BACKGROUND: Tidal marshes will be threatened by increasing rates of sea-level rise (SLR) over the next century. Managers seek guidance on whether existing and restored marshes will be resilient under a range of potential future conditions, and on prioritizing marsh restoration and conservation activities. METHODOLOGY: Building upon established models, we developed a hybrid approach that involves a mechanistic treatment of marsh accretion dynamics and incorporates spatial variation at a scale relevant for conservation and restoration decision-making. We applied this model to San Francisco Bay, using best-available elevation data and estimates of sediment supply and organic matter accumulation developed for 15 Bay subregions. Accretion models were run over 100 years for 70 combinations of starting elevation, mineral sediment, organic matter, and SLR assumptions. Results were applied spatially to evaluate eight Bay-wide climate change scenarios. PRINCIPAL FINDINGS: Model results indicated that under a high rate of SLR (1.65 m/century), short-term restoration of diked subtidal baylands to mid marsh elevations (-0.2 m MHHW) could be achieved over the next century with sediment concentrations greater than 200 mg/L. However, suspended sediment concentrations greater than 300 mg/L would be required for 100-year mid marsh sustainability (i.e., no elevation loss). Organic matter accumulation had minimal impacts on this threshold. Bay-wide projections of marsh habitat area varied substantially, depending primarily on SLR and sediment assumptions. Across all scenarios, however, the model projected a shift in the mix of intertidal habitats, with a loss of high marsh and gains in low marsh and mudflats. CONCLUSIONS/SIGNIFICANCE: Results suggest a bleak prognosis for long-term natural tidal marsh sustainability under a high-SLR scenario. To minimize marsh loss, we recommend conserving adjacent uplands for marsh migration, redistributing dredged sediment to raise elevations, and concentrating restoration efforts in sediment-rich areas. To assist land managers, we developed a web-based decision support tool (www.prbo.org/sfbayslr).

Niches, models, and climate change: assessing the assumptions and uncertainties.

As the rate and magnitude of climate change accelerate, understanding the consequences becomes increasingly important. Species distribution models (SDMs) based on current ecological niche constraints are used to project future species distributions. These models contain assumptions that add to the uncertainty in model projections stemming from the structure of the models, the algorithms used to translate niche associations into distributional probabilities, the quality and quantity of data, and mismatches between the scales of modeling and data. We illustrate the application of SDMs using two climate models and two distributional algorithms, together with information on distributional shifts in vegetation types, to project fine-scale future distributions of 60 California landbird species. Most species are projected to decrease in distribution by 2070. Changes in total species richness vary over the state, with large losses of species in some "hotspots" of vulnerability. Differences in distributional shifts among species will change species co-occurrences, creating spatial variation in similarities between current and future assemblages. We use these analyses to consider how assumptions can be addressed and uncertainties reduced. SDMs can provide a useful way to incorporate future conditions into conservation and management practices and decisions, but the uncertainties of model projections must be balanced with the risks of taking the wrong actions or the costs of inaction. Doing this will require that the sources and magnitudes of uncertainty are documented, and that conservationists and resource managers be willing to act despite the uncertainties. The alternative, of ignoring the future, is not an option.

Re-shuffling of species with climate disruption: a no-analog future for California birds?

By facilitating independent shifts in species' distributions, climate disruption may result in the rapid development of novel species assemblages that challenge the capacity of species to co-exist and adapt. We used a multivariate approach borrowed from paleoecology to quantify the potential change in California terrestrial breeding bird communities based on current and future species-distribution models for 60 focal species. Projections of future no-analog communities based on two climate models and two species-distribution-model algorithms indicate that by 2070 over half of California could be occupied by novel assemblages of bird species, implying the potential for dramatic community reshuffling and altered patterns of species interactions. The expected percentage of no-analog bird communities was dependent on the community scale examined, but consistent geographic patterns indicated several locations that are particularly likely to host novel bird communities in the future. These no-analog areas did not always coincide with areas of greatest projected species turnover. Efforts to conserve and manage biodiversity could be substantially improved by considering not just future changes in the distribution of individual species, but including the potential for unprecedented changes in community composition and unanticipated consequences of novel species assemblages.