Sex-specific recruitment rates contribute to male-biased sex ratio in Adélie penguins.

Sex-related differences in vital rates that drive population change reflect the basic life history of a species. However, for visually monomorphic bird species, determining the effect of sex on demographics can be a challenge. In this study, we investigated the effect of sex on apparent survival, recruitment, and breeding propensity in the Adélie penguin (Pygoscelis adeliae), a monochromatic, slightly size dimorphic species with known age, known sex, and known breeding history data collected during 1996-2019 (n = 2127 birds) from three breeding colonies on Ross Island, Antarctica. Using a multistate capture-mark-recapture maximum-likelihood model, we estimated apparent survival (S^), recapture (resighting) probability (p^), and the probability of transitioning among breeding states and moving between colonies (ψ^; colony-specific non-juvenile pre-breeders, breeders, and non-breeders). Survival rate varied by breeding status and colony, but not sex, and pre-breeders had higher survival rates than breeders and non-breeders. Females had a higher probability of recruiting into the breeding population each year and may enter the breeding pool at younger ages. In contrast, both sexes had the same probability of breeding from year to year once they had recruited. Although we detected no direct sex effects on survival, the variation in recruitment probability and age-at-first reproduction, along with lower survival rates of breeders compared to pre-breeders, likely leads to shorter lifespans for females. This is supported by our findings of a male-biased mean adult sex ratio (ASR) of 1.4 males for every female (x^ proportion of males = 0.57, SD = 0.07) across all colonies and years in this metapopulation. Our study illustrates how important it can be to disentangle sex-related variation in population vital rates, particularly for species with complex life histories and demographic dynamics.

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.

Genomes of Single-Stranded DNA Viruses in a Fecal Sample from South Polar Skua (Stercorarius maccormicki) on Ross Island, Antarctica.

South polar skuas migrate from subtropical regions to breed along coastal Antarctica. In a fecal sample collected on Ross Island, Antarctica, we identified 20 diverse microviruses (Microviridae) that share low levels of similarity to currently known microviruses; 6 appear to use a Mycoplasma/Spiroplasma codon translation table.

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.

Foraging dive frequency predicts body mass gain in the Adélie penguin.

Quantifying food intake in wild animals is crucial to many ecological and evolutionary questions, yet it can be very challenging, especially in the marine environment. Because foraging behavior can be inferred from dive recordings in many marine creatures, we hypothesized that specific behavioral dive variables can indicate food intake. To test this hypothesis, we attached time-depth recorders to breeding Adélie penguins also implanted with RFID tags that crossed a weighbridge as they traveled to and from the ocean to feed their chicks. The weighbridge reported how much mass the penguin had gained during a foraging trip. The variables that explained a significant amount of the change in body mass while at sea were the number of foraging dives per hour (46%) and the number of undulations per hour (12%). Most importantly, every increment of 1 in the rate of foraging dives per hour equated to a penguin gaining an average 170 g of mass, over the course of a 6-60 h foraging trip. These results add to a growing understanding that different metrics of foraging success are likely appropriate for different species, and that assessing the types and frequencies of dives using time-depth recorders can yield valuable insights.

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.

Age-related reproductive performance of the Adélie penguin, a long-lived seabird exhibiting similar outcomes regardless of individual life-history strategy.

Age-related variation in reproductive performance in long-lived iteroparous vertebrate species is common, with performance being influenced by within-individual processes, such as improvement and senescence, in combination with among-individual processes, such as selective appearance and disappearance. Few studies of age-related reproductive performance have compared the role of these drivers within a metapopulation, subject to varying degrees of resource competition. We accounted for within- and among-individual changes among known-aged Adélie penguins Pygoscelis adeliae during 17 years (1997-2013), at three clustered colonies of disparate size, to understand patterns in age-related reproductive success during early and late adulthood. Age at first reproduction (AFR) was lowest, and number of breeding attempts highest, at the largest colony. Regardless of AFR, success improved with early post-recruitment experience. For both oldest and youngest recruitment groups, peak performance occurred at the end of their reproductive life span indicating a possible cost of reproduction. Intermediate recruitment groups reached peak performance in their mid-reproductive life span and with intermediate breeding experience, before decreasing. Breeding success was lowest for the initial breeding attempt regardless of AFR, but we observed subsequent variation relative to recruitment age. Gaining experience by delaying recruitment positively influenced reproductive performance early in the reproductive life span and was most evident for the youngest breeders. Oldest recruits had the highest initial and peak breeding success. Differences in AFR resulted in trade-offs in reproductive life span or timing of senescence but not in the overall number of breeding attempts. Patterns differed as a function of colony size, and thus competition for resources. Early life improvement in performance at the larger colonies was primarily due to within-individual factors and at the largest colony, AFR. Regardless of colony size late-life performance was positively related to the age at last reproduction, indicating selective disappearance of lower performing individuals. These results highlight that different life-history strategies were equally successful, indicating that individuals can overcome potential trade-offs associated with early- and late-life performance. These results have important implications for understanding the evolution of life-history strategies responsible for driving population change.

Multidrone aerial surveys of penguin colonies in Antarctica.

Speed is essential in wildlife surveys due to the dynamic movement of animals throughout their environment and potentially extreme changes in weather. In this work, we present a multirobot path-planning method for conducting aerial surveys over large areas designed to make the best use of limited flight time. Unlike current survey path-planning solutions based on geometric patterns or integer programs, we solve a series of satisfiability modulo theory instances of increasing complexity. Each instance yields a set of feasible paths at each iteration and recovers the set of shortest paths after sufficient time. We implemented our planning algorithm with a team of drones to conduct multiple photographic aerial wildlife surveys of Cape Crozier, one of the largest Adélie penguin colonies in the world containing more than 300,000 nesting pairs. Over 2 square kilometers was surveyed in about 3 hours. In contrast, previous human-piloted single-drone surveys of the same colony required over 2 days to complete. Our method reduces survey time by limiting redundant travel while also allowing for safe recall of the drones at any time during the survey. Our approach can be applied to other domains, such as wildfire surveys in high-risk weather conditions or disaster response.

Drivers of concentrated predation in an Antarctic marginal-ice-zone food web.

Predators impact preyscapes (3-D distribution of forage species) by consuming prey according to their abilities or by altering prey behavior as they avoid being consumed. We elucidate prey (Antarctic silverfish[Pleuragramma antarctica] and crystal krill[Euphausia chrystallorophias]) responses to predation associated with the marginal ice zone (MIZ) of the McMurdo Sound, Antarctica, polynya. Prey abundance and habitat was sampled across a 30 × 15 km area by remotely-operated vehicle, and included locations that were accessible (ice edge) or inaccessible (solid fast ice) to air-breathing predators. Prey and habitat sampling coincided with bio-logging of Adélie penguins and observations of other air-breathing predators (penguins, seals, and whales), all of which were competing for the same prey. Adélie penguins dived deeper, and more frequently, near the ice edge. Lowered abundance of krill at the ice edge indicated they were depleted or were responding to increased predation and/or higher light levels along the ice edge. Penguin diet shifted increasingly to silverfish from krill during sampling, and was correlated with the arrival of krill-eating whales. Behaviorally-mediated, high trophic transfer characterizes the McMurdo Sound MIZ, and likely other MIZs, warranting more specific consideration in food web models and conservation efforts.

Tracking of marine predators to protect Southern Ocean ecosystems.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change1,2. Mitigation requires the identification and protection of Areas of Ecological Significance (AESs), which have so far not been determined at the ocean-basin scale. Here, using assemblage-level tracking of marine predators, we identify AESs for this globally important region and assess current threats and protection levels. Integration of more than 4,000 tracks from 17 bird and mammal species reveals AESs around sub-Antarctic islands in the Atlantic and Indian Oceans and over the Antarctic continental shelf. Fishing pressure is disproportionately concentrated inside AESs, and climate change over the next century is predicted to impose pressure on these areas, particularly around the Antarctic continent. At present, 7.1% of the ocean south of 40°S is under formal protection, including 29% of the total AESs. The establishment and regular revision of networks of protection that encompass AESs are needed to provide long-term mitigation of growing pressures on Southern Ocean ecosystems.

The retrospective analysis of Antarctic tracking data project.

The Retrospective Analysis of Antarctic Tracking Data (RAATD) is a Scientific Committee for Antarctic Research project led jointly by the Expert Groups on Birds and Marine Mammals and Antarctic Biodiversity Informatics, and endorsed by the Commission for the Conservation of Antarctic Marine Living Resources. RAATD consolidated tracking data for multiple species of Antarctic meso- and top-predators to identify Areas of Ecological Significance. These datasets and accompanying syntheses provide a greater understanding of fundamental ecosystem processes in the Southern Ocean, support modelling of predator distributions under future climate scenarios and create inputs that can be incorporated into decision making processes by management authorities. In this data paper, we present the compiled tracking data from research groups that have worked in the Antarctic since the 1990s. The data are publicly available through biodiversity.aq and the Ocean Biogeographic Information System. The archive includes tracking data from over 70 contributors across 12 national Antarctic programs, and includes data from 17 predator species, 4060 individual animals, and over 2.9 million observed locations.

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.

Evidence of age-related improvement in the foraging efficiency of Adélie penguins.

Age variation in reproductive performance is well-documented but the mechanisms underlying this variation remain unclear. Foraging efficiency is likely to be a key source of demographic variation as it determines the amount of energy that can be invested in fitness-related activities. Evidence of age-related changes in the foraging efficiency of adult seabirds is scarce and inconsistent. We investigated the effects of age on the foraging efficiency of breeding Adélie penguins, a relatively short-lived seabird species, in order to gain a broader perspective on the processes driving variation in ageing rates. We found support for a positive effect of age, either linear or levelling off at old ages, on both our proxies for daily catch rate and catch per unit effort. Across all age classes, males were more performant foragers than females. We found no strong evidence for differing ageing patterns between sexes or individual quality levels, and no evidence for senescence. We infer that continuous individual improvement could be responsible for a larger amount of the variation in foraging efficiency with age at our study site, compared with selective disappearance of underperforming phenotypes. The different results reported by other studies highlight the need to conduct longitudinal studies across a range of species in different environments.

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.

Diverse and highly recombinant anelloviruses associated with Weddell seals in Antarctica.

The viruses circulating among Antarctic wildlife remain largely unknown. In an effort to identify viruses associated with Weddell seals (Leptonychotes weddellii) inhabiting the Ross Sea, vaginal and nasal swabs, and faecal samples were collected between November 2014 and February 2015. In addition, a Weddell seal kidney and South Polar skua (Stercorarius maccormicki) faeces were opportunistically sampled. Using high throughput sequencing, we identified and recovered 152 anellovirus genomes that share 63-70% genome-wide identities with other pinniped anelloviruses. Genome-wide pairwise comparisons coupled with phylogenetic analysis revealed two novel anellovirus species, tentatively named torque teno Leptonychotes weddellii virus (TTLwV) -1 and -2. TTLwV-1 (n = 133, genomes encompassing 40 genotypes) is highly recombinant, whereas TTLwV-2 (n = 19, genomes encompassing three genotypes) is relatively less recombinant. This study documents ubiquitous TTLwVs among Weddell seals in Antarctica with frequent co-infection by multiple genotypes, however, the role these anelloviruses play in seal health remains unknown.

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.

Circumpolar analysis of the Adélie Penguin reveals the importance of environmental variability in phenological mismatch.

Evidence of climate-change-driven shifts in plant and animal phenology have raised concerns that certain trophic interactions may be increasingly mismatched in time, resulting in declines in reproductive success. Given the constraints imposed by extreme seasonality at high latitudes and the rapid shifts in phenology seen in the Arctic, we would also expect Antarctic species to be highly vulnerable to climate-change-driven phenological mismatches with their environment. However, few studies have assessed the impacts of phenological change in Antarctica. Using the largest database of phytoplankton phenology, sea-ice phenology, and Adélie Penguin breeding phenology and breeding success assembled to date, we find that, while a temporal match between Penguin breeding phenology and optimal environmental conditions sets an upper limit on breeding success, only a weak relationship to the mean exists. Despite previous work suggesting that divergent trends in Adélie Penguin breeding phenology are apparent across the Antarctic continent, we find no such trends. Furthermore, we find no trend in the magnitude of phenological mismatch, suggesting that mismatch is driven by interannual variability in environmental conditions rather than climate-change-driven trends, as observed in other systems. We propose several criteria necessary for a species to experience a strong climate-change-driven phenological mismatch, of which several may be violated by this system.

Sex-Based Differences in Adélie Penguin (Pygoscelis adeliae) Chick Growth Rates and Diet.

Sexually size-dimorphic species must show some difference between the sexes in growth rate and/or length of growing period. Such differences in growth parameters can cause the sexes to be impacted by environmental variability in different ways, and understanding these differences allows a better understanding of patterns in productivity between individuals and populations. We investigated differences in growth rate and diet between male and female Adélie Penguin (Pygoscelis adeliae) chicks during two breeding seasons at Cape Crozier, Ross Island, Antarctica. Adélie Penguins are a slightly dimorphic species, with adult males averaging larger than adult females in mass (~11%) as well as bill (~8%) and flipper length (~3%). We measured mass and length of flipper, bill, tibiotarsus, and foot at 5-day intervals for 45 male and 40 female individually-marked chicks. Chick sex was molecularly determined from feathers. We used linear mixed effects models to estimate daily growth rate as a function of chick sex, while controlling for hatching order, brood size, year, and potential variation in breeding quality between pairs of parents. Accounting for season and hatching order, male chicks gained mass an average of 15.6 g d(-1) faster than females. Similarly, growth in bill length was faster for males, and the calculated bill size difference at fledging was similar to that observed in adults. There was no evidence for sex-based differences in growth of other morphological features. Adélie diet at Ross Island is composed almost entirely of two species--one krill (Euphausia crystallorophias) and one fish (Pleuragramma antarctica), with fish having a higher caloric value. Using isotopic analyses of feather samples, we also determined that male chicks were fed a higher proportion of fish than female chicks. The related differences in provisioning and growth rates of male and female offspring provides a greater understanding of the ways in which ecological factors may impact the two sexes differently.

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.