Regional genetic structure in the Magellanic penguin (Spheniscus magellanicus) suggests metapopulation dynamics

Using microsatellite markers and mitochondrial DNA (mtDNA) sequences (cytochrome oxidase 1 gene), we estimated levels of genetic structuring among nine Magellanic Penguin (Spheniscus magellanicus) colonies distributed throughout three major reproductive regions of the South Atlantic Ocean. Overall, breeding colonies showed relatively high levels of genetic diversity at both nuclear and mtDNA markers (mean heteorzygosity: He = 0.598; mean allelic diversity: A = 7.11; mtDNA haplotype diversity: h = 0.812). A hierarchical analysis of molecular variance based on microsatellite data showed limited genetic structuring of breeding colonies, with 99% of the variation explained by differences among individuals and 0.7–1.0% attributed to differences among the three regions. The mtDNA analysis revealed higher levels of genetic structuring, with 3.43% of the variation explained by regions and 2.24% explained by colonies within the regions. Furthermore, a Mantel test revealed a significant association between geographic and genetic distances among colonies. The limited genetic structuring we detected is likely a result of (1) population intermixing through natal dispersal and (2) the large effective sizes of the reproductive colonies, both of which prevent genetic differentiation at neutral markers, balanced with (3) the regional association of breeding colonies to distinct feeding grounds and (4) a recent expansion of the population. Our results suggest that the demographic dynamics of breeding colonies of Magellanic Penguins may be framed under a metapopulation model, in which colonies with large numbers of breeding pairs could be considered source populations for maintaining the overall abundance of this species in the Atlantic Ocean

Physiological condition in Magellanic Penguins: Does it matter if you have to walk a long way to your nest?

Colony edges, as opposed to interiors, are often considered less advantageous nesting places in colonial species. For temperate-breeding penguins, inland colony edges should be less desirable than other edges, as there are added costs of walking farther inland, and ambient temperatures are higher. During settlement and incubation, we compared body condition and baseline and stress-induced levels of the hormone corticosterone in male Magellanic Penguins (Spheniscus magellanicus) nesting on the sea edge of a colony with those nesting on the inland edge, \u3e800 m from shore. Body condition in both groups was significantly lower during settlement than during incubation, but was similar in both groups within breeding stages. Corticosterone levels were similar between breeding stages and for groups within each breeding stage. While body condition can vary over time, penguins appear to be well buffered to physiological extremes, as they do not show modification of corticosterone levels with variations in nesting conditions

Sex Ratio is Variable and Increasingly Male Biased at Two Colonies of Magellanic Penguins

Sex ratios are commonly skewed and variable in wild populations, but few studies track temporal trends in this demographic parameter. We examined variation in the operational sex ratio at two protected and declining breeding colonies of Magellanic Penguins (Spheniscus magellanicus) in Chubut, Argentina. Penguins from the two colonies, separated by 105 km, migrate north in the non‐breeding season and have overlapping distributions at sea. Conditions during the non‐breeding season can impact long‐term trends in operational sex ratio (i.e., through sex‐specific survival) and interannual variation in operational sex ratio (i.e., through sex‐specific breeding decisions). We found an increasingly male‐biased operational sex ratio at the two disparate colonies of Magellanic Penguins, which may contribute to continued population decline. We also found that the two colonies showed synchronous interannual variation in operational sex ratio, driven by variation in the number of females present each year. This pattern may be linked to sex‐specific overwintering effects that cause females to skip breeding, i.e., to remain at sea rather than returning to the colony to breed, more often than males. Contrary to our predictions, colony‐wide reproductive success was not lower in years with a more male‐biased operational sex ratio. We did find that males showed more evidence of fighting and were less likely to pair when the operational sex ratio was more male biased. Our results highlight an indirect mechanism through which variation in the operational sex ratio can influence populations, through a higher incidence of fighting among the less abundant sex. Because biased sex ratios can reduce the size of the breeding population and influence rates of conflict, tracking operational sex ratio is critical for conservation

Age and food deprivation affects expression of the glucocorticosteriod stress response in Magellanic penguin (Spheniscus magellanicus) chicks

We examined how the glucocortical stress response in free‐living Magellanic penguin (Spheniscus magellanicus) chicks changes with age and whether adrenocortical function of chicks within a brood varies in relation to food provisioned by adults. Chicks showed little corticosterone response to capture stress shortly after hatching, an intermediate response around 45‐d posthatch, and a robust stress response near fledging. However, in response to an adrenocorticotropic hormone (ACTH) challenge, hatchlings were capable of secreting corticosterone at adult‐like levels. The larger sibling in broods of two showed a similar gradual stress‐response development pattern. In contrast, by day 45, when differences in body condition were well established between siblings, the smaller, food‐deprived chicks significantly increased baseline levels of corticosterone but showed normal stress‐induced levels. Near fledging, baseline levels had returned to normal, but stress‐induced levels were lower than expected. Similar to altricial species, normally developing semialtricial Magellanic penguin chicks do not express a robust corticosterone stress response until near fledging. Chronic stressors such as food deprivation cause corticosterone use to be up‐regulated earlier than expected. However, in cases of extended chronic stress, down‐regulation may ensue, thus avoiding the negative effects of chronically elevated levels of corticosterone

The Role of Demographic History and Selection in Shaping Genetic Diversity of the Galápagos Penguin (Spheniscus mendiculus)

Although many studies have documented the effects of demographic bottlenecks on the genetic diversity of natural populations, there is conflicting evidence of the roles that genetic drift and selection may play in driving changes in genetic variation at adaptive loci. We analyzed genetic variation at microsatellite and mitochondrial loci in conjunction with an adaptive MHC class II locus in the Galápagos penguin (Spheniscus mendiculus), a species that has undergone serial demographic bottlenecks associated with El Niño events through its evolutionary history. We compared levels of variation in the Galápagos penguin to those of its congener, the Magellanic penguin (Spheniscus magellanicus), which has consistently maintained a large population size and thus was used as a non-bottlenecked control. The comparison of neutral and adaptive markers in these two demographically distinct species allowed assessment of the potential role of balancing selection in maintaining levels of MHC variation during bottleneck events. Our analysis suggests that the lack of genetic diversity at both neutral and adaptive loci in the Galápagos penguin likely resulted from its restricted range, relatively low abundance, and history of demographic bottlenecks. The Galápagos penguin revealed two MHC alleles, one mitochondrial haplotype, and six alleles across five microsatellite loci, which represents only a small fraction of the diversity detected in Magellanic penguins. Despite the decreased genetic diversity in the Galápagos penguin, results revealed signals of balancing selection at the MHC, which suggest that selection can mitigate some of the effects of genetic drift during bottleneck events. Although Galápagos penguin populations have persisted for a long time, increased frequency of El Niño events due to global climate change, as well as the low diversity exhibited at immunological loci, may put this species at further risk of extinction

Population regulation in Magellanic penguins: what determines changes in colony size?

Seabirds are often studied at individual colonies, but the confounding effects of emigration and mortality processes in open populations may lead to inappropriate conclusions on the mechanisms underlying population changes. Magellanic penguin (Spheniscus magellanicus) colonies of variable population sizes are distributed along the Argentine coastline. In recent decades, several population and distributional changes have occurred, with some colonies declining and others newly established or increasing. We integrated data of eight colonies scattered along ~600 km in Northern Patagonia (from 41°26´S, 65°01´W to 45°11´S, 66°30´W, Rio Negro and Chubut provinces) and conducted analysis in terms of their growth rates, production of young and of the dependence of those vital rates on colony age, size, and location. We contrasted population trends estimated from abundance data with those derived from population modeling to understand if observed growth rates were attainable under closed population scenarios. Population trends were inversely related to colony size, suggesting a density dependent growth pattern. All colonies located in the north — which were established during the last decades — increased at high rates, with the smallest, recently established colonies growing at the fastest rate. In central-southern Chubut, where colonies are the oldest, the largest breeding aggregations declined, but smaller colonies remained relatively stable. Results provided strong evidence that dispersal played a major role in driving local trends. Breeding success was higher in northern colonies, likely mediated by favorable oceanographic conditions. However, mean foraging distance and body condition of chicks at fledging were influenced by colony size. Recruitment of penguins in the northern area may have been triggered by a combination of density dependence, likely exacerbated by less favorable oceanographic conditions in the southern sector. Our results reaffirm the idea that individual colony trends do not provide confident indicators of population health, highlighting the need to redefine the scale for the study of population changes.Fil: Pozzi, Luciana Melina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Nacional Patagónico; ArgentinaFil: Garcia Borboroglu, Jorge Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Nacional Patagónico; Argentina. Global Penguin Society. Washington; Estados Unidos. University of Washington; Estados UnidosFil: Boersma, P. Dee. Global Penguin Society. Washington; Estados Unidos. University of Washington; Estados UnidosFil: Pascual, Miguel Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Nacional Patagónico; Argentina. Universidad Nacional de la Patagonia; Argentin

Lateralization (handedness) in Magellanic penguins

Lateralization, or asymmetry in form and/or function, is found in many animal species. Brain lateralization is considered adaptive for an individual, and often results in “handedness,” “footedness,” or a side preference, manifest in behavior and morphology. We tested for lateralization in several behaviors in a wild population of Magellanic penguins Spheniscus magellanicus breeding at Punta Tombo, Argentina. We found no preferred foot in the population (each penguin observed once) in stepping up onto an obstacle: 53% stepped up with the right foot, 47% with the left foot (n = 300, binomial test p = 0.27). We found mixed evidence for a dominant foot when a penguin extended a foot for thermoregulation, possibly depending on the ambient temperature (each penguin observed once). Penguins extended the right foot twice as often as the left foot (n = 121, p < 0.0005) in 2 years when we concentrated our effort during the heat of the day. In a third year when we observed penguins early and late in the day, there was no preference (n = 232, p = 0.59). Penguins use their flippers for swimming, including searching for and chasing prey. We found morphological evidence of a dominant flipper in individual adults: 60.5% of sternum keels curved one direction or the other (n = 76 sterna from carcasses), and 11% of penguins had more feather wear on one flipper than the other (n = 1217). Right-flippered and left-flippered penguins were equally likely in both samples (keels: p = 0.88, feather wear: p = 0.26), indicating individual but not population lateralization. In fights, aggressive penguins used their left eyes preferentially, consistent with the right side of the brain controlling aggression. Penguins that recently fought (each penguin observed once) were twice as likely to have blood only on the right side of the face (69%) as only on the left side (31%, n = 175, p < 0.001). The proportion of penguins with blood only on the right side increased with the amount of blood. In most fights, the more aggressive penguin used its left eye and attacked the other penguin’s right side. Lateralization depended on the behavior tested and, in thermoregulation, likely on the temperature. We found no lateralization or mixed results in the population of Magellanic penguins in three individual behaviors, stepping up, swimming, and thermoregulation. We found lateralization in the population in the social behavior fighting

Happy Feet in a Hostile World? The Future of Penguins Depends on Proactive Management of Current and Expected Threats

Penguins face a wide range of threats. Most observed population changes have been negative and have happened over the last 60 years. Today, populations of 11 penguin species are decreasing. Here we present a review that synthesizes details of threats faced by the world’s 18 species of penguins. We discuss alterations to their environment at both breeding sites on land and at sea where they forage. The major drivers of change appear to be climate, and food web alterations by marine fisheries. In addition, we also consider other critical and/or emerging threats, namely human disturbance near nesting sites, pollution due to oil, plastics and chemicals such as mercury and persistent organic compounds. Finally, we assess the importance of emerging pathogens and diseases on the health of penguins. We suggest that in the context of climate change, habitat degradation, introduced exotic species and resource competition with fisheries, successful conservation outcomes will require new and unprecedented levels of science and advocacy. Successful conservation stories of penguin species across their geographical range have occurred where there has been concerted effort across local, national and international boundaries to implement effective conservation planning

Occurrence of Magellanic Penguins along the Northeast Brazilian Coast during 2008 Austral Winter

During the austral winter of 2008, thousands of penguins traveled to low latitudes along the South Atlantic coast of South America. The atmospheric and oceanic conditions from April to July 2008 may account for the penguins' unusual geographic distribution. During that period, South Atlantic coastal waters were cooler; the wind anomalies had northward and onshore components; the ocean's coastal region presented northward currents that favored the penguins to travel toward lower latitudes. This anomalous climate regime resulted from extreme meteorological frontal systems that occurred mainly during June 2008. Three consecutive extreme midlatitude cyclones produced strong wind shear that resulted in the northward oceanic flow along the South American eastern shoreline favoring the penguins to be spotted in northern tropical waters

Linking multidimensional functional diversity to quantitative methods: a graphical hypothesis-evaluation framework

Functional trait analysis is an appealing approach to study differences among biological communities because traits determine species' responses to the environment and their impacts on ecosystem functioning. Despite a rapidly expanding quantitative literature, it remains challenging to conceptualize concurrent changes in multiple trait dimensions (“trait space”) and select quantitative functional diversity methods to test hypotheses prior to analysis. To address this need, we present a widely applicable framework for visualizing ecological phenomena in trait space to guide the selection, application, and interpretation of quantitative functional diversity methods. We describe five hypotheses that represent general patterns of responses to disturbance in functional community ecology and then apply a formal decision process to determine appropriate quantitative methods to test ecological hypotheses. As a part of this process, we devise a new statistical approach to test for functional turnover among communities. Our combination of hypotheses and metrics can be applied broadly to address ecological questions across a range of systems and study designs. We illustrate the framework with a case study of disturbance in freshwater communities. This hypothesis-driven approach will increase the rigor and transparency of applied functional trait studies.Keywords: community assembly, trait-based ecology, disturbance, ordination, functional diversity, multivariate analysis, multidimensional trait spaceThis is the publisher’s final pdf. The article is copyrighted by Ecological Society of America and published by John Wiley & Sons, Inc. It can be found at: http://esajournals.onlinelibrary.wiley.com/hub/journal/10.1002/%28ISSN%291939-9170