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Pododermatitis is common in penguins kept under human care. Substrate optimization plays an important role in prevention and treatment; however, there is limited information on biomechanical properties of commonly used substrates on penguin feet. The objectives were to test the ability of different substrates to decrease weight loading on the central metatarsal pad of penguin feet in an ex vivo model using feet with and without bumblefoot harvested from two Magellanic penguin ( Spheniscus magellanicus ) cadavers. Penguin feet were attached to a digital force gauge mounted onto a stand for compression testing at 2.5 and 5 kg. Forces at the central metatarsal pad were measured in triplicate using small force sensors. Tested substrates included five granular surfaces (sand, wet sand, pea gravel, wet pea gravel, and crushed ice), three compliant surfaces (short-leaf Astroturf, long-leaf Astroturf, and neoprene), and three firm surfaces (tile, rubber drainage mat, and 3M Safety-Walk Wet Area Matting). Data were analyzed using linear mixed models. There were multifaceted effects of applied pressures, substrate surfaces, and pododermatitis on central metatarsal measured pressures. In general, doubling compression forces resulted in higher measured pressures in all firm and compliant surfaces but not in granular surfaces. Firm surfaces were associated with higher recorded plantar pressures at 2.5 kg, but different significance groupings emerged at 5 kg with a high-, medium-, and low-pressure cluster of surfaces. Pododermatitis lesions resulted in significant alterations in statistical significance clustering among substrate surfaces and unique substrate behaviors. The results of this study could help in making recommendations pertaining to foot health for penguin exhibits.

The illustration is an important tool to aid in the description and understanding of anatomy, and penguins (Aves: Sphenisciformes: Spheniscidae) are an important clade in environmental monitoring, paleontology, and other research fields. Traditionally, anatomic illustration has been informed by dissection. More recently, micro-computed tomography (micro-CT) has proven to be a powerful tool for three-dimensional anatomic imaging, although larger specimens are more challenging to image due to increased X-ray attenuation. Here, we used traditional dissection and micro-CT to illustrate the skulls of Aptenodytes patagonicus, Eudyptula minor, and Pygoscelis papua, and the extracranial soft tissue of E. minor. Micro-CT prevented the loss of orientation, disarticulation, and distortion of bones that might result from cleaning and drying skulls, while immobilization was achieved by freezing the specimens before imaging. All bony elements in the head were accurately depicted. Fixing, dehydrating, and diffusion staining with iodine (diceCT) enabled the identification of muscles and other large nonmineralized structures, but specimen preparation precluded the ability to show smaller nerves and vessels. The results presented here provide a guide for anatomic studies of penguins and our summary of sample preparation and imaging techniques are applicable for studies of other similarly sized biological specimens., (© 2022 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

In order to evaluate mercury (Hg) accumulation patterns in Southern Ocean penguins, we measured Hg concentrations and carbon (δ 13 C) and nitrogen (δ 15 N) stable isotope ratios in body feathers of adult Adélie (Pygoscelis adeliae), gentoo (Pygoscelis papua), and chinstrap (Pygoscelis antarctica) penguins living near Anvers Island, West Antarctic Peninsula (WAP) collected in the 2010/2011 austral summer. With these and data from Pygoscelis and other penguin genera (Eudyptes and Aptenodytes) throughout the Southern Ocean, we modelled Hg variation using δ 13 C and δ 15 N values. Mean concentrations of Hg in feathers of Adélie (0.09 ± 0.05 μg g -1 ) and gentoo (0.16 ± 0.08 μg g -1 ) penguins from Anvers Island were among the lowest ever reported for the Southern Ocean. However, Hg concentrations in chinstrap penguins (0.80 ± 0.20 μg g -1 ), which undertake relatively broad longitudinal winter migrations north of expanding sea ice, were significantly higher (P < 0.001) than those in gentoo or Adélie penguins. δ 13 C and δ 15 N values for feathers from all three Anvers Island populations were also the lowest among those previously reported for Southern Ocean penguins foraging within Antarctic and subantarctic waters. These observations, along with size distributions of WAP krill, suggest foraging during non-breeding seasons as a primary contributor to higher Hg accumulation in chinstraps relative to other sympatric Pygoscelis along the WAP. δ 13 C values for all Southern Ocean penguin populations, alone best explained feather Hg concentrations among possible generalized linear models (GLMs) for populations grouped by either breeding site (AICc = 36.9, w i  = 0.0590) or Antarctic Frontal Zone (AICc = 36.9, w i  = 0.0537). Although Hg feather concentrations can vary locally by species, there was an insignificant species-level effect (w i  < 0.001) across the full latitudinal range examined. Therefore, feeding ecology at breeding locations, as tracked by δ 13 C, control Hg accumulation in penguin populations across the Southern Ocean., Competing Interests: Declaration of competing interest The authors declare no competing financial interests., (Copyright © 2024 Elsevier B.V. All rights reserved.)

The prevalence of microplastics (MPs, <5 mm) in natural environments presents a formidable global environmental threat MPs can be found from the Arctic to Antarctica, including glaciers. Despite their widespread distribution, studies on MP accumulation in apex predators inhabiting Polar Regions remain limited. The objective of this study was to conduct a comprehensive examination, for the first time, of MP bioaccumulation in various organs and tissue of Adélie penguins. This investigation comprehends the gastrointestinal tract (GIT), scat, internal organ (lung, trachea, spleen, and liver) and tissue (muscle) samples collected from Svenner Island, Antarctica during the 39th Indian expedition to Antarctica in 2019-2020. Our analyses revealed the presence of 34 MPs across the GIT, scat, lung, and trachea samples, with no MPs detected in muscle, spleen, or liver tissues. Blue-colored microfibers (>50 %) and MPs smaller than 1 mm (38 %) in size were prominently observed. Polymer characterization utilizing μ-FTIR spectroscopy identified low-density polyethylene (LDPE) (~63 %) as the predominant polymer type. The accumulation of MP fibers in the gastrointestinal tract and scat of Adélie penguins may originate from marine ambient media and prey organisms. Furthermore, the presence of LDPE fibers in the trachea and lungs likely occurred through inhalation and subsequent deposition of MPs originating from both local and long-range airborne sources. The identification of fibers ranging between 20 and 100 μm within the trachea suggests a plausible chance of cellular deposition of MPs. Overall our findings provide valuable insights into the organ-specific accumulation of MPs in apex predators. Adélie penguins emerge as promising environmental bio-monitoring species, offering insights into the potential trophic transfer of MPs within frigid environments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Satellite-based remote sensing and uncrewed aerial imagery play increasingly important roles in the mapping of wildlife populations and wildlife habitat, but the availability of imagery has been limited in remote areas. At the same time, ecotourism is a rapidly growing industry and can yield a vast catalog of photographs that could be harnessed for monitoring purposes, but the inherently ad-hoc and unstructured nature of these images make them difficult to use. To help address this, a subfield of computer vision known as phototourism has been developed to leverage a diverse collection of unstructured photographs to reconstruct a georeferenced three-dimensional scene capturing the environment at that location. Here we demonstrate the use of phototourism in an application involving Antarctic penguins, sentinel species whose dynamics are closely tracked as a measure of ecosystem functioning, and introduce a semi-automated pipeline for aligning and registering ground photographs using a digital elevation model (DEM) and satellite imagery. We employ the Segment Anything Model (SAM) for the interactive identification and segmentation of penguin colonies in these photographs. By creating a textured 3D mesh from the DEM and satellite imagery, we estimate camera poses to align ground photographs with the mesh and register the segmented penguin colony area to the mesh, achieving a detailed representation of the colony. Our approach has demonstrated promising performance, though challenges persist due to variations in image quality and the dynamic nature of natural landscapes. Nevertheless, our method offers a straightforward and effective tool for the georegistration of ad-hoc photographs in natural landscapes, with additional applications such as monitoring glacial retreat., Competing Interests: The authors have declared that no competing interests exist., (Copyright: This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.)

The proportions of individuals in various age classes in a population of wild animals affect population trends, behaviors, learning, and social structures. Knowledge of age structure is needed for effective conservation and management of populations. However, it is not always possible to determine the age or age class of individual animals, and hence the age structure of the population. Penguins, like most birds, cannot be aged once they molt into adult plumage. Spheniscus penguins attain adult plumage at 6 to 24 months of age, and individuals can live more than 30 years. We studied foot darkening in the four species of Spheniscus penguins to determine if age class can be determined from foot color. We compared how foot color changes with age among the four species to investigate potential functions of the darkening. We found that Spheniscus penguins have pale feet at hatching and the feet become darker with age throughout the lives of individuals. We showed that we can accurately predict the age structure of a colony of Magellanic penguins Spheniscus magellanicus , but not the ages of individual penguins, based on a sample of foot colors. The timing of foot darkening within species was consistent with foot color functioning in protection from UV radiation, and not with foot color functioning in thermoregulation. The species that breeds at the lowest latitudes and experiences the highest UV radiation (Galápagos penguins Spheniscus mendiculus ) had feet that darkened at the earliest ages, and the species that breed at higher latitudes and experience less intense insolation (African S. demersus and Magellanic penguins) had feet that darkened latest. Humboldt penguins S. humboldti breed mostly at low latitudes and foot darkening was intermediate between Galápagos and Magellanic penguins. We also found that males' feet darken somewhat earlier than females' feet, likely because females spend more time in their nests (burrows or under vegetation) than males and have less sun exposure. We found that feet darkened in an individual over years, but not within a breeding season. The color change is a life-long process, likely an evolutionary adaptation within species, not a seasonal, temporary response to UV radiation. We propose foot darkening as a way to assess age structure in Spheniscus penguins. Foot color in a colony of Magellanic penguins can provide a rapid, noninvasive method to estimate the age structure of the colony., Competing Interests: Ginger A. Rebstock and P. Dee Boersma are Academic Editors for PeerJ., (©2024 Rebstock et al.)

The present work extends the scope of prior studies through analysis, modelling and simulation of the As, Cd, Co, Cu, Fe Mn, Mo, Ni and Zn release from Gentoo (Pygoscelis papua) and Chinstrap (Pygoscelis antarcticus) penguin guano to the Southern Ocean seawater and to Antarctic snow meltwater. Laboratory experimental results have been modelled considering kinetic processes between water and guano using two element pools in the guano compartment; its application allows us to interpret behaviours and predict release concentrations of dissolved trace elements from guano which are potentially useful for incorporation as elements source into biogeochemical models applied in the Southern Ocean. Variations in quantities and release patterns depending on the type of guano and aqueous medium in contact have been identified. The release percentages from the guano to the aqueous medium, once the steady state has been reached, vary depending on the water medium and guano type in the ranges of 100-2.9 % for Mo; 91.5-68.6 % for Ni; 81.8-22.8 % As; 52.0-43.9 % Cu; 26.9-7.4 % Mn; 24.9-5.4 for Co; 4.4-3.2 % for Zn and 0.94-0.51 % for Fe. Considering a penguin population of 774,000 Gentoo and 8,000,000 Chinstrap, the estimated annual mass released to the both seawater and freshwater would be ≈18,500 kg for Cu, ≈1710 kg for Zn, ≈1944 kg for Fe, ≈1640 kg for Mn, ≈499 kg for As, ≈289 kg for Ni, ≈155 kg for Mo, ≈36.7 kg for Cd and ≈8.1 kg for Co. These contributions can be locally significant both in promoting phytoplankton growth and in their role as inhibitors of primary productivity., Competing Interests: Declaration of competing interest The authors declare that they have no know competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Our research focuses on semen collection from 42 African penguin males, grouped by age, habituation levels, and reproductive season timing. We assess behavioral and physiological responses to dorso-abdominal massage, evaluating species-specific seminal traits using conventional and advanced methods. Positive behavioral responses corresponded with successful semen collection. Young and human-reared males exhibited more favorable behaviors, and samples containing spermatozoa were more likely collected during the reproductive season. Age did not influence sample collectability or spermatozoa presence, though mature males produced higher semen volumes. Young males exhibited more sperm morphological abnormalities, especially outside the breeding season. Sperm concentration and number per ejaculate showed no significant differences across age or seasonal groups. Young males had higher motile and progressive sperm percentages, while mature males had more static sperm. Additionally, percentages of live sperm and intact acrosomes were higher in mature males. Young males and samples from non-reproductive season presented more dead sperm with damaged acrosomes. Plasma membrane integrity positively correlated with age for live sperm and negatively for dying sperm. This research demonstrates the feasibility of semen collection from African penguins regardless of habituation level, fills the knowledge gap by describing sperm morphological abnormalities, and provides insights into using flow cytometry in Sphenisciformes., (© 2024. The Author(s).)

Rationale: Understanding the migration of marine animals is hindered by the limitations of traditional tracking methods. It is therefore crucial to develop alternative methods. Stable isotope-based tracking has proven useful for this task, although it requires detailed isoscapes in the focal area. Here, we present predator-based isoscapes of the coastal zone of the Patagonian Shelf Large Marine Ecosystem (PSLME), which offers a novel tool for geolocation., Methods: Whole-blood samples from breeding Magellanic penguins nesting at 11 colonies were used to create δ 15 N and δ 13 C isoscapes. Isotopic values were assigned to random positions inside their corresponding foraging area. Spatial analysis and data interpolation resulted in δ 15 N and δ 13 C isoscapes for the coastal zone of the PSLME, which were validated through cross-validation., Results: The isoscapes mean standard error ranged from 0.05 to 0.41 for δ 15 N and from 0.07 to 0.3 for δ 13 C, similar to the error range of the mass spectrometer used for measuring isotope ratios. Predictive surfaces reflected the latitudinal trends, with δ 13 C and δ 15 N values increasing northwards. δ 13 C values showed a strong latitudinal gradient, while δ 15 N values had two distinct domains, with higher values in the north. The error surface indicated the highest certainty within 130 km from the shore and within the reported Magellanic penguin foraging areas., Conclusions: Both isoscapes revealed strong spatial variation. The δ 13 C isoscape showed a latitudinal gradient, consistent with patterns in other oceans. The δ 15 N isoscape clearly separated northern and southern colonies, likely influenced by nitrogen sources. The error obtained fell within the measurement error ranges, adding credibility to the models., (© 2024 John Wiley & Sons Ltd.)

The Antarctic krill Euphausia superba is a key species in Antarctic waters, mainly because it is a major component of the diet of dominant predators, including penguins. It is also a marine living resource that is commercially harvested. Since 2000, krill fishing has become more concentrated. On the basis of several years of data, it has been reported that up to 57% of the krill fishery harvests occur in the Bransfield Strait area. The distribution of krill in the Southern Ocean is not well described. Studies that compare the size of krill caught by commercial vessels with that recorded in the diet of predators are even rarer. The main objectives of this study were to assess the spatial diversity in the size and putative age of krill in the CCAMLR subarea 48.1, to investigate the spatial availability of krill and its size and age structure, and to assess whether the diet of Pygoscelis penguins reflects the size structure of krill present in the environment. The results implied that the size and age structure of the krill population were similar throughout the Bransfield Strait during the study period, although those in the eastern and southern parts of the strait and the Brabant Island region were the most similar. The Livingston Island and Drake Passage areas were clearly distinguishable from the above regions, where larger and therefore older krill were recorded. All Pygoscelis penguin species showed size preferences for consumed krill; therefore, their diet is likely not a reliable indicator of the size of krill in the environment. Krill that had not yet reached sexual maturity, and thus not yet started reproducing were commercially caught in the Bransfield Strait during the investigated years., (© 2024. The Author(s).)

Byssus is a unique external structure in sessile bivalves and is critical for settlement and metamorphosis. However, little is known about the stout byssus in Pteria penguin. We explored the byssus structure and proteins using scanning electron microscopy and proteomics, respectively. The results revealed that P. penguin byssus has a dense and highly aligned fiber inner core, and the outer cuticle contains protein granules embedded in the protein matrix. Proteomic analysis revealed 31 proteins in the byssus, among which 15 differentially expressed proteins were mainly enriched in the EGF/EGF-like and laminin EGF-like domains. Foot proteins were enriched in the EF-hand, immunoglobulin, and fibronectin domains. All these domains can participate in protein-protein and/or protein-metal interactions in the extracellular matrix (ECM), which, together with the seven types of ECM proteins detected in the byssus, supports the hypothesis that the byssus is derived from the ECM. We also found that in vitro acellular structures of the byssus and the shell shared commonalities in their formation processes. These results are useful for further understanding byssus evolution and the characterization of byssus-related proteins. SIGNIFICANCE: This manuscript investigates the structure and the origin of Pteria penguin byssus, given that byssus is vital to provide critical protection for reproduction and even against environmental stresses that affect survival. However, there is rare research on byssus protein composition. Hence, though scanning electron microscopy and proteomic analysis, we discovered that P. penguin byssus possesses the dense and highly aligned fiber inner core, and the outer cuticle has protein granules embedded in the protein matrix. Proteomic analysis showed that there were 31 proteins in the byssus, among which 15 proteins were mainly enriched in the EGF/EGF-like and laminin EGF-like domains. Foot proteins closely related to byssus formation were enriched in EF hand, immunoglobulin, and fibronectin domains. These domains are able to participate in protein-protein and/or protein-metal interactions in the extracellular matrix (ECM), which together with the seven types of ECM proteins detected in byssus support the hypothesis that byssus derive from the ECM. We also found in vitro acellular structures the byssus and the shell share commonalities in their formation processes. These results were useful for further understanding the byssus evolution and the characterization of the byssus-related proteins., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Fecal corticosterone metabolite (FCM) concentrations, which can be determined noninvasively, have recently been explored as a stress indicator in birds. In our study, we measured FCM concentrations in penguins under nonmolting or molting conditions, cool or hot season, diseased condition, and incubation period. These measurements were conducted in an aquarium that housed king penguins, gentoo penguins, and African penguins. This study aimed to investigate the validity of fecal matter as a stress indicator. Our findings revealed that FCM concentrations were significantly higher in molting individuals than in nonmolting individuals. Compared with the cool season, FCM concentrations were significantly higher in penguins housed outdoors during the hot season. However, no differences were observed in penguins housed indoors. Diseased individuals and an incubating individual showed notably higher FCM concentrations than healthy individuals. Interestingly, the FCM concentration in king penguin that underwent cataract surgery was extremely high before the surgery. However, 1 month postsurgery, it decreased to a level similar to that of healthy individuals. We observed increased FCM concentrations in penguins considered to be exposed to stressors. Notably, FCM concentration decreased after removing the stress factor. The FCM concentration was found to be consistent with the stress state of penguins, suggesting its usefulness as a stress indicator.

The relative importance of genetic drift and local adaptation in facilitating speciation remains unclear. This is particularly true for seabirds, which can disperse over large geographic distances, providing opportunities for intermittent gene flow among distant colonies that span the temperature and salinity gradients of the oceans. Here, we delve into the genomic basis of adaptation and speciation of banded penguins, Galápagos (Spheniscus mendiculus), Humboldt (Spheniscus humboldti), Magellanic (Spheniscus magellanicus), and African penguins (Spheniscus demersus), by analyzing 114 genomes from the main 16 breeding colonies. We aim to identify the molecular mechanism and genomic adaptive traits that have facilitated their diversifications. Through positive selection and gene family expansion analyses, we identified candidate genes that may be related to reproductive isolation processes mediated by ecological thermal niche divergence. We recover signals of positive selection on key loci associated with spermatogenesis, especially during the recent peripatric divergence of the Galápagos penguin from the Humboldt penguin. High temperatures in tropical habitats may have favored selection on loci associated with spermatogenesis to maintain sperm viability, leading to reproductive isolation among young species. Our results suggest that genome-wide selection on loci associated with molecular pathways that underpin thermoregulation, osmoregulation, hypoxia, and social behavior appears to have been crucial in local adaptation of banded penguins. Overall, these results contribute to our understanding of how the complexity of biotic, but especially abiotic, factors, along with the high dispersal capabilities of these marine species, may promote both neutral and adaptive lineage divergence even in the presence of gene flow., (© The Author(s) 2024. Published by Oxford University Press on behalf of Society for Molecular Biology and Evolution.)

The gut microbiomes of Antarctic penguins are important for the fitness of the host birds and their chicks. The compositions of microbial communities in Antarctic penguin guts are strongly associated with the birds' diet, physiological adaptation and phylogeny. Whilst seasonal changes in food resources, distribution and population parameters of Antarctic penguins have been well addressed, little research is available on the stability or variability of penguin stomach microbiomes over time. Here, we focused on two Pygoscelis penguin species breeding sympatrically in the maritime Antarctic and analysed their stomach contents to assess whether penguin gut microbiota differed over three austral summer breeding seasons. We used a high-throughput DNA sequencing approach to study bacterial diversity in stomach regurgitates of Adélie ( Pygoscelis adeliae ) and chinstrap ( Pygoscelis antarctica ) penguins that have a similar foraging regime on Signy Island (South Orkney Islands). Our data revealed significant differences in bacterial alpha and beta diversity between the study seasons. We also identified bacterial genera that were significantly associated with specific breeding seasons, diet compositions, chick-rearing stages and sampling events. This study provides a baseline for establishing future monitoring of penguin gut microbiomes in a rapidly changing environment.

Antemortem serodiagnosis of aspergillosis remains challenging in Sphenisciformes. Protein electrophoresis, serology (antibody, antigen) by ELISA, and gliotoxin detection provide variable diagnostic value. In the present study, a commercially available Western blot (WB) validated for use in humans and dolphins was adapted for use with penguin samples. Using the same method and reagents, samples were analyzed from multiple institutions in the United States and one facility in France. This was inclusive of normal juvenile African penguins ( Spheniscus demersus , n = 10) and various species of penguins in the United States with confirmed infection (n = 9) as well as 52 samples from Humboldt penguins ( Spheniscus humboldti ) in France. Cumulative WB scores (based on reactivity to different antigens) were found to be significantly higher in the group of penguins with confirmed infection (p < 0.0001). Significant differences were also observed between the clinically normal penguins in the two populations, with higher scores in the United States (median score 1.0, 95%CI [0-5], min 0, max 11) compared to France (median score 0,95%CI [0-0], min 0, max 5). The utilization of the WB as a diagnostic tool is inconclusive due to the use of samples from varying institutions, environmental background, age, and stages of infection. However, this tool may provide an overview of antigen reactivity in penguins infected with Aspergillus to help design a more robust serology assay and further understand the humoral immune response during infection.

Environmental microbes routinely colonize wildlife body surface microbiota. However, animals experience dynamic environmental shifts throughout their daily routine. Yet, the effect of ecological shifts in wildlife body surface microbiota has been poorly explored. Here, we sequenced the hypervariable region V3-V4 of the 16S rRNA gene to characterize the body surface microbiota of wild Magellanic penguins (Spheniscus magellanicus) under two ecological contexts: (1) Penguins walking along the coast and (2) Penguins sheltered underground in their nest, across three subantarctic breeding colonies in the Magellan Strait, Chile. Despite ecological contexts, our results revealed that Moraxellaceae bacteria were the most predominant and abundant taxa associated with penguin body surfaces. Nevertheless, we detected colony-specific core bacteria associated with penguin bodies. The most abundant were: Deinococcus in the Contramaestre colony, Fusobacterium in the Tuckers 1 colony, and Clostridium sensu stricto 1 in the Tuckers 2 colony. Our results give a new perspective on the niche environmental hypothesis for wild seabirds. First, the ecological characteristics of each colony were associated with the microbial communities from the nest soil and the body surface of penguins inside the nests. For example, in the colonies with heterogenous vegetation cover (i.e. the Tuckers Islets), there was a similar microbial composition between the nest soil and the body surface of penguins. In contrast, on the more arid colony (Contramaestre), we detected differences in the microbial communities between the nest soil and the body surface of penguins., (© 2024 The Author(s). Molecular Ecology published by John Wiley & Sons Ltd.)

The African penguin is currently experiencing a significant decline, with just over 10,000 breeding pairs left. A substantial body of research reflects the impacts of contaminants of emerging concern (CECs) on the marine environment, with wastewater treatment plants reported as one of the main sources of CEC release. In South Africa, CECs were identified contaminating the marine environment and bioaccumulating in several marine species. Approximately 70 % of all African penguin colonies breed in close proximity to cities and/or harbors in South Africa. Currently, the impact of CECs as a stressor upon the viability of African penguin populations is unknown. Based on the search results there was a clear lack of information on CECs' bioaccumulation and impact on the African penguin. This narrative review will thus focus on the prevalent sources and types of CECs and examine the reported consequences of constant exposure in seabirds, particularly African penguins., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

A multitude of animal species undergo prolonged fasting events at regularly occurring life history stages. During such periods of food deprivation, individuals need to suppress their appetite. The satiety signalling gut hormone ghrelin has received much attention in this context in studies looking at mammalian systems. In wild birds, however, knowledge on the ghrelin system and its role during extended fasts is still scarce. In this study, we collected plasma samples for measurements of circulating ghrelin concentrations from adult southern rockhopper penguins (Eudyptes chrysocome chrysocome) during the three to four week-long moult-fast that they repeat annually to replace their feathers. We further sampled chicks before and after feeding bouts and non-moulting adults. Circulating ghrelin levels did not differ significantly between fed and unfed chicks but chicks had significantly lower plasma ghrelin levels compared to adults. Furthermore, penguins in late moult (i.e. individuals at the end of the prolonged fasting bout) had higher ghrelin levels compared to non-moulting adults. Our results show elevated levels of circulating ghrelin during moult and generally lower levels of ghrelin in chicks than in adults regardless of feeding state. Given the scarcity or absence of knowledge on the function of ghrelin in seabirds and in fasting birds in general, our results add greatly to our understanding of the avian ghrelin system., Competing Interests: Declaration of competing interest None., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Aspergillosis is a major cause of morbidity and mortality in penguins, with triazole antifungal drugs being commonly used for prophylaxis and treatment. This report describes 15 cases of fatal hemolysis associated with liquid itraconazole and voriconazole formulations administered to African penguins ( Spheniscus demersus ) from four institutions. All penguins underwent stressful events (e.g. relocation, induced molt) and were administered commercial liquid itraconazole formulations or compounded voriconazole liquid suspension. Observed clinical signs in affected penguins prior to death included hyporexia, weight loss, lethargy, dyspnea, red-tinged droppings, and obtunded mentation. Intra- and extravascular hemolysis and hemoglobinuric nephrosis were the primary pathologic manifestations on postmortem examination. The concentration-dependent hemolytic potentials of itraconazole, voriconazole, and commercial and compounded vehicle suspensions were evaluated in vitro by exposing chicken whole blood as a surrogate for penguin blood. Hemoglobin content in blood plasma was then measured by spectrophotometry. Neither itraconazole nor voriconazole alone induced hemolysis in vitro. The vehicle ingredients sorbitol and hydromellose induced hemolysis, but not at predicted plasma levels in chicken erythrocytes, suggesting neither the azole antifungals nor their major vehicles alone were likely to contribute to hemolysis in vivo in these penguins. Potential mechanisms of toxicosis include generation of an unmeasured reactive metabolite causing hemolysis, preexisting erythrocyte fragility, or species-specific differences in hemolytic thresholds that were not assessed in the chicken erythrocyte model. More research is needed on the potential for toxicosis of azole antifungal drugs and carrier molecules in this and other avian species.

B-cell lymphoma/leukemia-2 (BCL2), an anti-apoptotic factor in the mitochondrial regulatory pathway of apoptosis, is critically important in immune defenses. In this study, a novel BCL2 gene was characterized from Pteria penguin (P. penguin). The PpBCL2 was 1482 bp long, containing an open reading frame (ORF) of 588 bp encoding 195 amino acids. Four highly conserved BCL-2 homology (BH) domains were found in PpBCL2. Amino acid alignment and phylogenetic tree showed that PpBCL2 had the highest similarity with BCL2 of Crassostrea gigas at 65.24 %. Tissue expression analysis showed that PpBCL2 had high constitutive expression in gill, digestive diverticulum and mantle, and was significantly increased 72 h of Vibrio parahaemolyticus (V. parahaemolyticus) challenge in these immune tissues. Furthermore, PpBCL2 silencing significantly inhibited antimicrobial activity of hemolymph supernatant by 1.4-fold, and significantly reduced the survival rate by 51.7 % at 72 h post infection in P. penguin. These data indicated that PpBCL2 played an important role in immune response of P. penguin against V. parahaemolyticus infection., Competing Interests: Declaration of competing interest The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Evolutionary rescue, whereby adaptive evolutionary change rescues populations from extinction, is theorized to enable imperiled animal populations to persist under increasing anthropogenic change. Despite a large body of evidence in theoretical and laboratory settings, the potential for evolutionary rescue to be a viable adaptation process for free-ranging animals remains unknown. Here, we leverage a 38-year dataset following the fates of 53,959 Magellanic penguins ( Spheniscus magellanicus ) to investigate whether a free-ranging vertebrate species can morphologically adapt to long-term environmental change sufficiently to promote population persistence. Despite strong selective pressures, we found that penguins did not adapt morphologically to long-term environmental changes, leading to projected population extirpation. Fluctuating selection benefited larger penguins in some environmental contexts, and smaller penguins in others, ultimately mitigating their ability to adapt under increasing environmental variability. Under future climate projections, we found that the species cannot be rescued by adaptation, suggesting similar constraints for other long-lived species. Such results reveal how fluctuating selection driven by environmental variability can inhibit adaptation under long-term environmental change. Our eco-evolutionary approach helps explain the lack of adaptation and evolutionary rescue in response to environmental change observed in many animal species., Competing Interests: Competing interests statement:The authors declare no competing interest.

Numerous penguins can propagate pathogens with antibiotic resistance genes (ARGs) into Antarctica. However, the effects of penguin dissemination on the lake ARGs still have received little attention via guano deposition. Here, we have profiled ARGs in ornithogenic sediments subject to penguin guano (OLS) and nonornithogenic sediments (NOLS) from 16 lakes across Antarctica. A total of 191 ARGs were detected in all sediment samples, with a much higher abundance and diversity in OLS than in NOLS. Surprisingly, highly diverse and abundant ARGs were found in the OLS with a detection frequency of >40% and an absolute abundance of (2.34 × 10 9 )-(4.98 × 10 9 ) copies g -1 , comparable to those in coastal estuarine sediments and pig farms. The strong correlations of identified resistance genes with penguin guano input amount, environmental factors, mobile genetic elements, and bacterial community, in conjunction with network and redundancy analyses, all indicated that penguins were responsible for the dissemination and high enrichment of ARGs in lake sediments via the guano deposition, which might greatly outweigh local human-activity effects. Our results revealed that ARGs could be carried into lakes across the Antarctica through penguin migration, food chains, and guano deposition, which were closely connected with the widespread pollution of ARGs at the global scale.

The microbiome is a key factor in the health, well-being, and success of vertebrates, contributing to the adaptive capacity of the host. However, the impact of geographic and biotic factors that may affect the microbiome of wild birds in polar environments is not well defined. To address this, we determined the bacterial 16S rRNA gene sequence profiles in faecal samples from pygoscelid penguin populations in the Scotia Arc, focusing on gentoo penguins. This mesopredatory group breeds in defined colonies across a wide geographic range. Since diet could influence microbiome structure, we extracted dietary profiles from a eukaryotic 18S rRNA gene sequence profile. The bacterial microbiome profiles were considered in the context of a diverse set of environmental and ecological measures. Integrating wide geographic sampling with bacterial 16S and eukaryotic 18S rRNA gene sequencing of over 350 faecal samples identified associations between the microbiome profile and a suite of geographic and ecological factors. Microbiome profiles differed according to host species, colony identity, distance between colonies, and diet. Interestingly there was also a relationship between the proportion of host DNA (in relation to total 18S rRNA gene signal) and the microbiome, which may reflect gut passage time. Colony identity provided the strongest association with differences in microbiome profiles indicating that local factors play a key role in the microbiome structure of these polar seabirds. This may reflect the influence of local transfer of microbes either via faecal-oral routes, during chick feeding or other close contact events. Other factors including diet and host species also associate with variation in microbiome profile, and in at least some locations, the microbiome composition varies considerably between individuals. Given the variation in penguin microbiomes associated with diverse factors there is potential for disruption of microbiome associations at a local scale that could influence host health, productivity, and immunological competence. The microbiome represents a sensitive indicator of changing conditions, and the implications of any changes need to be considered in the wider context of environmental change and other stressors., (© 2024. The Author(s).)

Phenotypic differences often stem from genetic/maternal differences and/or early-life adaptations to local environmental conditions. In colonial animals, little is known on how variation in the social environment is embedded into individual phenotypes, nor what the consequences are on individual fitness. We conducted an experimental cross-fostering study on king penguins ( Aptenodytes patagonicus ), exchanging eggs among 134 pairs breeding in high-density (67 pairs) or low-density (67 pairs) areas of the same breeding colony. We investigated differences in parent and chick phenotypes and survival in relation to the density of their origin and foster environment. Adults breeding in colony areas of high density exhibited decreased resting behaviour and increased aggression and vigilance, increased hypometabolism during incubation fasts, and more moderate corticosterone responses shaped by exposure to chronic stressors (e.g. constant aggression by neighbours). Chick phenotypes were more influenced by the environment in which they were raised than their genetic/maternal origin. Chicks raised in high-density colonial environments showed enhanced weight gain and survival rates regardless of the density of their genetic parents' breeding areas. Our study experimentally shows advantages to breeding in colonial areas of higher breeder densities in king penguins, and highlights the importance of social settings in shaping phenotype expression in colonial seabirds.

Understanding how populations respond to variability in environmental conditions and interspecific interactions is one of the biggest challenges of population ecology, particularly in the context of global change. Although many studies have investigated population responses to climate change, very few have explicitly integrated interspecific relationships when studying these responses. In this study, we aimed to understand the combined effects of interspecific interactions and environmental conditions on the demographic parameters of a prey-predator system of three sympatric seabird populations breeding in Antarctica: the south polar skua (Catharacta maccormicki) and its two main preys during the breeding season, the Adélie penguin (Pygoscelis adeliae) and the emperor penguin (Aptenodytes forsteri). We built a two-species integrated population model (IPM) with 31 years of capture-recapture and count data and provided a framework that made it possible to estimate the demographic parameters and abundance of a predator-prey system in a context where capture-recapture data were not available for one species. Our results showed that predator-prey interactions and local environmental conditions differentially affected south polar skuas depending on their breeding state of the previous year. Concerning prey-predator relationships, the number of Adélie penguin breeding pairs showed a positive effect on south polar skua survival and breeding probability, and the number of emperor penguin dead chicks showed a positive effect on the breeding success of south polar skuas. In contrast, there was no evidence for an effect of the number of south polar skuas on the demography of Adélie penguins. We also found an important impact of sea ice conditions on both the dynamics of south polar skuas and Adélie penguins. Our results suggest that this prey-predator system is mostly driven by bottom-up processes and local environmental conditions., (© 2024 The Author(s). Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Estimating the densities of marine prey observed in animal-borne video loggers when encountered by foraging predators represents an important challenge for understanding predator-prey interactions in the marine environment. We used video images collected during the foraging trip of one chinstrap penguin (Pygoscelis antarcticus) from Cape Shirreff, Livingston Island, Antarctica to develop a novel approach for estimating the density of Antarctic krill (Euphausia superba) encountered during foraging activities. Using the open-source Video and Image Analytics for a Marine Environment (VIAME), we trained a neural network model to identify video frames containing krill. Our image classifier has an overall accuracy of 73%, with a positive predictive value of 83% for prediction of frames containing krill. We then developed a method to estimate the volume of water imaged, thus the density (N·m-3) of krill, in the 2-dimensional images. The method is based on the maximum range from the camera where krill remain visibly resolvable and assumes that mean krill length is known, and that the distribution of orientation angles of krill is uniform. From 1,932 images identified as containing krill, we manually identified a subset of 124 images from across the video record that contained resolvable and unresolvable krill necessary to estimate the resolvable range and imaged volume for the video sensor. Krill swarm density encountered by the penguins ranged from 2 to 307 krill·m-3 and mean density of krill was 48 krill·m-3 (sd = 61 krill·m-3). Mean krill biomass density was 25 g·m-3. Our frame-level image classifier model and krill density estimation method provide a new approach to efficiently process video-logger data and estimate krill density from 2D imagery, providing key information on prey aggregations that may affect predator foraging performance. The approach should be directly applicable to other marine predators feeding on aggregations of prey., Competing Interests: The authors have declared that no competing interests exist., (Copyright: This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.)

A 21-year-old captive female Humboldt penguin (Spheniscus humboldti) was in good health until the day before the animal died. However, the animal suddenly exhibited symptoms of vomiting and anorexia, and died in shortly thereafter. The autopsy revealed a blood clot in the abdominal air sac and 280 mL of dark red fluid in the body cavity. The ovary was twisted around a blood vessel. Based on the findings described, it was considered that the cause of death was hypovolemic shock due to massive intracoelomic hemorrhage by vascular collapse resulting from ovarian torsion. This is the first report of ovarian torsion in penguins.

Microplastic pollution threatens some of the world's most iconic locations for marine biodiversity, including the remote Galápagos Islands, Ecuador. Using the Galápagos penguin (Spheniscus mendiculus) as a sentinel species, the present study assessed microplastics and suspected anthropogenic cellulose concentrations in surface seawater and zooplankton near Santa Cruz and Galápagos penguin colonies (Floreana, Isabela, Santiago), as well as in penguin potential prey (anchovies, mullets, milkfish) and penguin scat. On average, 0.40 ± 0.32 microplastics L -1 were found in surface seawater (<10 μm; n = 63 L), while 0.003, 0.27, and 5.12 microplastics individual -1 were found in zooplankton (n = 3372), anchovies (n = 11), and mullets (n = 6), respectively. The highest concentration (27 microplastics individual -1 ) was observed in a single milkfish. Calculations based on microplastics per gram of prey, in a potential diet composition scenario, suggest that the Galápagos penguin may consume 2881 to 9602 microplastics daily from prey. Despite this, no microplastics or cellulose were identified in 3.40 g of guano collected from two penguins. Our study confirms microplastic exposure in the pelagic food web and endangered penguin species within the UNESCO World Heritage site Galápagos Islands, which can be used to inform regional and international policies to mitigate plastic pollution and conserve biodiversity in the global ocean. Environ Toxicol Chem 2024;43:1442-1457. © 2024 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC., (© 2024 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.)

The aim of this study was to determine the minimum anesthetic concentration of isoflurane (MAC ISO ) and sevoflurane (MAC SEVO ) and evaluate the cardiorespiratory changes induced by varying fractions of inspired oxygen (FiO 2 ) in Magellanic penguins (Spheniscus magellanicus). Twenty adult penguins (3.53 ± 0.44 kg) of undetermined sex were used. Both MAC ISO (n = 9) and MAC SEVO (n = 13) were established using an up-and-down design. Next, twelve mechanically ventilated penguins were maintained at 1 MAC ISO or 1 MAC SEVO (n = 6 per group) with the FiO 2 initially set at 1.0. Three FiO 2 values (0.6, 0.4 and 0.2) were then held constant during anesthesia for 20 minutes each. Arterial blood samples were collected for gas analysis after the 20-minute period for each FiO 2 . Mean ± SD MAC ISO was 1.93 ± 0.10% and MAC SEVO was 3.53 ± 0.13%. Other than heart rate at 0.6 FiO 2 (86 ± 11 beats/minute in MAC ISO and 132 ± 37 beats/minute in MAC SEVO ; p = 0.041), no significant cardiorespiratory differences were detected between groups. In both groups, decreasing the FiO 2 produced increased pH values and reduced partial pressures of carbon dioxide and bicarbonate. Partial pressures of oxygen (PaO 2 ) gradually lowered from 1.0 FiO 2 through 0.2 FiO 2 , though hypoxemia (PaO 2  < 80 mmHg) occurred only with the latter FiO 2 . The MAC ISO and the MAC SEVO for the Magellanic penguin fell within the upper range of reported avian MAC estimates. To prevent hypoxemia in healthy, mechanically ventilated, either isoflurane- or sevoflurane-anesthetized Magellanic penguins, a minimum FiO 2 of 0.4 should be used., (© 2024. The Author(s), under exclusive licence to Springer Nature B.V.)

The highly pathogenic Avian Influenza virus (HPAIV) H5N1 has caused a global outbreak affecting both wild and domestic animals, predominantly avian species. To date, cases of the HPAIV H5 Clade 2.3.4.4b in penguins have exclusively been reported in African Penguins. In Chile, the virus was confirmed in pelicans in December 2022 and subsequently spread across the country, affecting several species, including Humboldt penguins. This study aims to provide an overview of the incidents involving stranded and deceased Humboldt penguins and establish a connection between these events and HPAIV H5N1. Historical data about strandings between 2009 and 2023 was collected, and samples from suspected cases in 2023 were obtained to confirm the presence of HPAIV H5N1. Between January and August 2023, 2,788 cases of stranded and deceased penguins were recorded. Out of these, a total of 2,712 penguins deceased, evidencing a significative increase in mortality starting in early 2023 coinciding with the introduction and spreading of HPAIV H5N1 in the country. Thirty-seven events were categorized as mass mortality events, with the number of deceased penguins varying from 11 to 98. Most cases (97 %) were observed in the North of Chile. One hundred and eighty-one specimens were subjected to HPAIV diagnosis, four of which tested positive for HPAIV H5N1. Spatial analysis validates the correlation between mass mortality events and outbreaks of HPAIV in Chile. However, the limited rate of HPAIV H5N1 detection, which can be attributed to the type and quality of the samples, requiring further exploration., Competing Interests: Declaration of Competing Interest The authors declare no conflict of interest., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Once a year, penguins undergo a catastrophic moult, replacing their entire plumage during a fasting period on land or on sea-ice during which time individuals can lose 45% of their body mass. In penguins, new feather synthesis precedes the loss of old feathers, leading to an accumulation of two feather layers (double coat) before the old plumage is shed. We hypothesized that the combination of the high metabolism required for new feather synthesis and the potentially high thermal insulation linked to the double coat could lead to a thermal challenge requiring additional peripheral circulation to thermal windows to dissipate the extra heat. To test this hypothesis, we measured the surface temperature of different body regions of captive gentoo penguins (Pygoscelis papua) throughout the moult under constant environmental conditions. The surface temperature of the main body trunk decreased during the initial stages of the moult, suggesting greater thermal insulation. In contrast, the periorbital region, a potential proxy of core temperature in birds, increased during these same early moulting stages. The surface temperature of the bill, flipper and foot (thermal windows) tended to initially increase during the moult, highlighting the likely need for extra heat dissipation in moulting penguins. These results raise questions regarding the thermoregulatory capacities of penguins in the wild during the challenging period of moulting on land in the current context of global warming., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Aspergillus fumigatus is a common opportunistic pathogen in different animals, including birds such as penguins. For the first time, a fungal strain identified as A. fumigatus was isolated from soil in the nests of gentoo penguins, Pygoscelis papua, on Livingston Island, South Shetland Islands (maritime Antarctica). This isolate (A. fumigatus UFMGCB 11829) displayed a series of potentially pathogenic characteristics in vitro. We evaluated its detailed molecular taxonomy and submitted the A. fumigatus UFMGCB 11829 Antarctic strain to in vivo pathogenic modelling. The isolate was confirmed to represent A. fumigatus morphological and phylogenetic analysis showed that it was closely related to A. fumigatus sequences reported from animals, immunosuppressed humans, storage grains, plants and soils. The strain displayed the best mycelial growth and conidia production at 37 ºC; however, it was also able to grow and produce conidia at 15º, demonstrating its capability to survive and colonize penguin nest at least in the summer season in maritime Antarctica. In pathogenicity tests, healthy mice did not showed symptoms of infection; however, 50% lethality was observed in immunosuppressed mice that were inoculated with 10 6 and 10 7 spores. Lethality increased to 100% when inoculated with 10 8 spores. Our data highlight the potential pathogenicity of opportunistic A. fumigatus that may be present in the Antarctic, and the risks of both their further transfer within Antarctica and outwards to other continents, risks which may be exacerbated due global climatic changes., (© 2024. The Author(s) under exclusive licence to Sociedade Brasileira de Microbiologia.)

Scleral ossicles and other bony elements are present in the eyes of many vertebrates, including birds. In this study, the skeletal elements present in the penguin eye and orbit were imaged using macro photographs and micro-computed tomography (micro-CT), to help elucidate their function and significance. A total of 36 scleral rings and three whole skulls were imaged. King (Aptenodytes patagonicus), Fiordland crested (Eudyptes pachyrhynchus), Snares crested (Eudyptes robustus), royal (Eudyptes schlegeli) and yellow-eyed (Megadyptes antipodes) penguins had between 12 and 14 elements in their scleral ring while the gentoo (Pygoscelis papua) had 14 and 17; little penguins (Eudyptula sp.) consistently had between 10 and 12 elements. All had at least two elements that overlapped, usually totally, each neighbour, and two that were overlapped by each neighbour. The interior structure of all ossicles revealed a lattice-like arrangement of struts typical of cancellous bone, the whole being surrounded by thick cortical bone. The scleral ring of a 10 week gentoo chick was not completely ossified but rather had multiple small holes within it on micro-CT. A large os opticus was present in one king penguin but in another bird of the same age and gender there was no such bone. Much smaller accessory bones were found in the posterior pole of one Snares crested and one little penguin. We conclude that the penguin scleral ring not only maintains the shape of the eye but also provides protection and a site of insertion for rectus muscles. However, the extreme variability in the os opticus suggests that it is not essential to normal function., (© 2021 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

We report the discovery of a dead Magellanic penguin (Spheniscus magellanicus) found on Juquehy Beach (23°46'S 45°44'W), municipality of São Sebastião, Brazil, on September 9th 2020. Following necropsy, we noted the presence of an adult size PFF-2 protective mask within the stomach of the penguin which we inferred as the cause of death. As far as we are aware, this is the first recorded instance of marine animal mortality by protective face mask ingestion. Whilst concerns have been raised relating microplastic contamination in marine environments from Covid-19 related waste, there has been relatively less attention paid to the potential risk of macro-scale contaminants, such as protective face coverings. We suggest that Covid-19 related macro contaminants be considered in coastal marine risk assessments and urge further research on this topic., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Assessing the physiological stress responses of wild animals opens a window for understanding how organisms cope with environmental challenges. Since stress response is associated with changes in body temperature, the use of body surface temperature through thermal imaging could help to measure acute and chronic stress responses non-invasively. We used thermal imaging, acute handling-stress protocol and an experimental manipulation of corticosterone (the main glucocorticoid hormone in birds) levels in breeding king penguins (Aptenodytes patagonicus), to assess: 1. The potential contribution of the Hypothalamo-Pituitary-Adrenal (HPA) axis in mediating chronic and acute stress-induced changes in adult surface temperature, 2. The influence of HPA axis manipulation on parental investment through thermal imaging of eggs and brooded chicks, and 3. The impact of parental treatment on offspring thermal's response to acute handling. Maximum eye temperature (T eye ) increased and minimum beak temperature (T beak ) decreased in response to handling stress in adults, but neither basal nor stress-induced surface temperatures were significantly affected by corticosterone implant. While egg temperature was not significantly influenced by parental treatment, we found a surprising pattern for chicks: chicks brooded by the (non-implanted) partner of corticosterone-implanted individuals exhibited higher surface temperature (both T eye and T beak ) than those brooded by glucocorticoid-implanted or control parents. Chick's response to handling in terms of surface temperature was characterized by a drop in both T eye and T beak independently of parental treatment. We conclude that the HPA axis seems unlikely to play a major role in determining chronic or acute changes in surface temperature in king penguins. Changes in surface temperature may primarily be mediated by the Sympathetic-Adrenal-Medullary (SAM) axis in response to stressful situations. Our experiment did not reveal a direct impact of parental HPA axis manipulation on parental investment (egg or chick temperature), but a potential influence on the partner's brooding behaviour., Competing Interests: Declaration of competing interest Antoine Stier was an Associate Editor of J Thermal Biol at the time of submitting the manuscript but took no part in the peer review and decision-making processes for this research article., (Copyright © 2024 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Emperor penguins (Aptenodytes forsteri) are under increasing environmental pressure. Monitoring colony size and population trends of this Antarctic seabird relies primarily on satellite imagery recorded near the end of the breeding season, when light conditions levels are sufficient to capture images, but colony occupancy is highly variable. To correct population estimates for this variability, we develop a phenological model that can predict the number of breeding pairs and fledging chicks, as well as key phenological events such as arrival, hatching and foraging times, from as few as six data points from a single season. The ability to extrapolate occupancy from sparse data makes the model particularly useful for monitoring remotely sensed animal colonies where ground-based population estimates are rare or unavailable., (© 2024. The Author(s).)

Penguins are proficient swimmers, and their survival depends on their ability to catch prey. The diving behaviour of these fascinating birds should then minimize the associated energy cost. For the first time, the energy cost of penguin dives is computed from the free-ranging dive data, on the basis of an existing biomechanical model. Time-resolved acceleration and depth data collected for 300 dives of little penguins (Eudyptula minor) are specifically employed to compute the bird dive angles and swimming speeds, which are needed for the energy estimate. We find that the numerically obtained energy cost by using the free-ranging dive data is not far from the minimum cost predicted by the model. The outcome, therefore, supports the physical soundness of the chosen model; however, it also suggests that, for closer agreement, one should consider previously neglected effects, such as those due to water currents and those associated with motion unsteadiness. Additionally, from the free-ranging dive data, we calculate hydrodynamic forces and non-dimensional indicators of propulsion performance - Strouhal and Reynolds numbers. The obtained values further confirm that little penguins employ efficient propulsion mechanisms, in agreement with previous investigations., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Recent studies have highlighted the presence of antibiotic resistance genes (ARGs) in Antarctica, which are typically indicative of human activity. However, these studies have concentrated in the Antarctic Peninsula region, and relatively less is known about ARG prevalence in East Antarctica, where human activity levels are lower compared to the Antarctic Peninsula. In addition, the mechanisms of ARG transmission to Antarctica through natural or anthropogenic pathways remain unclear. In this study, we analyzed the fecal samples of Adélie penguins and South polar skuas by using high-throughput sequencing and microfluidic quantitative PCR to detect potential pathogens and ARGs at their breeding colonies near Syowa Station in East Antarctica. These results revealed the presence of several potential pathogens in the fecal matter of both bird species. However, the HF183 marker, which indicates human fecal contamination, was absent in all samples, as well as seawater sampled near the breeding colonies. This suggests that the human fecal contamination was negligible in our study area. In addition to pathogens, we found a significant number of ARGs and metal resistance genes in the feces of both Adélie penguins and South polar skuas, with higher detection rates in skuas than in penguins. To better understand how these birds acquire and transmit these genes, we analyzed the migratory patterns of Adélie penguins and South polar skuas by geolocator tracking. We found that the skuas migrate to the tropical and subtropical regions of the Indian Ocean during the austral winter. On the other hand, Adélie penguins exhibited a more localized migration pattern, mainly staying within Antarctic waters. Because the Indian Ocean is considered one of the major reservoirs of ARGs, South polar skuas might be exposed to ARGs during their winter migration and transfer these genes to Antarctica., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Group-living animals, including penguins, exhibit affiliative behaviors such as grooming (preening) and proximity. Such behaviors in non-primate animals have been less studied than those in primates. Our research focused on 20 identifiable Humboldt penguins in a zoo, analyzing kin relationships and reciprocity in preening and proximity by employing a 5-minute scan sampling method to observe and record individual behavior. Our findings revealed that preening and proximity were more prevalent among mate pairs. However, among non-mate pairs, such behaviors were more commonly observed between siblings and parent-offspring pairs. Notably, the individuals preened on each other simultaneously in all instances. This study highlights the potential influence of kin selection in shaping the affiliative behavior of penguins. Additionally, our findings indicate that penguins gain benefits from mutual preening. This study contributes to our understanding of social behaviors in non-primate species and emphasizes the need for further comparative studies of various animal taxa to elucidate the evolution of sociality., Competing Interests: Declaration of Competing Interest The authors declare no competing interests., (Copyright © 2024 Elsevier B.V. All rights reserved.)

How Antarctic species are facing historical and new stressors remains under-surveyed and risks to wildlife are still largely unknown. Adélie penguins Pygoscelis adeliae are well-known bioindicators and sentinels of Antarctic ecosystem changes, a true canary in the coal mine. Immuno-haematological parameters have been proved to detect stress in wild animals, given their rapid physiological response that allows them tracking environmental changes and thus inferring habitat quality. Here, we investigated variation in Erythrocyte Nuclear Abnormalities (ENAs) and White Blood Cells (WBCs) in penguins from three clustered colonies in the Ross Sea, evaluating immuno-haematological parameters according to geography, breeding stage, and individual penguin characteristics such as sex, body condition and nest quality. Concentrations of mercury (Hg) and stable isotopes of carbon and nitrogen (as proxies of the penguin's trophic ecology) were analysed in feathers to investigate the association between stress biomarkers and Hg contamination in Adélie penguins. Colony and breeding stage were not supported as predictors of immuno-haematological parameters. ENAs and WBCs were respectively ∼30 % and ∼20 % higher in male than in female penguins. Body condition influenced WBCs, with penguins in the best condition having a ∼22 % higher level of WBCs than those in the worst condition. Nest position affected the proportion of micronuclei (MNs), with inner-nesting penguins having more than three times the proportion of MNs than penguins nesting in peripheral positions. Heterophils:Lymphocytes (H:L) ratio was not affected by any of the above predictors. Multiple factors acting as stressors are expected to increase prominently in Antarctic wildlife in the near future, therefore extensive monitoring aimed to assess the health status of penguin populations is mandatory., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Marine pollution is becoming ubiquitous in the environment. Observations of pollution on beaches, in the coastal ocean, and in organisms in the Antarctic are becoming distressingly common. Increasing human activity, growing tourism, and an expanding krill fishing industry along the West Antarctic Peninsula all represent potential sources of plastic pollution and other debris (collectively referred to as debris) to the region. However, the sources of these pollutants from point (pollutants released from discrete sources) versus non-point (pollutants from a large area rather than a specific source) sources are poorly understood. We used buoyant simulated particles released in a high-resolution physical ocean model to quantify pollutant loads throughout the region. We considered non-point sources of debris from the Antarctic Circumpolar Current, Bellingshausen Sea, Weddell Sea, and point source pollution from human activities including tourism, research, and fishing. We also determined possible origins for observed debris based on data from the Southern Ocean Observing System and Palmer Long-Term Ecological Research program. Our results indicate that point source pollution released in the coastal Antarctic is more likely to serve as a source for observed debris than non-point sources, and that the dominant source of pollution is region-specific. Penguin colonies in the South Shetland and Elephant Islands had the greatest debris load from point sources whereas loads from non-point sources were greatest around the southernmost colonies. Penguin colonies at Cornwallis Island and Fort Point were exposed to the highest theoretical debris loads. While these results do not include physical processes such as windage and Stokes Drift that are known to impact debris distributions and transport in the coastal ocean, these results provide critical insights to building an effective stratified sampling and monitoring effort to better understand debris distributions, concentrations, and origins throughout the West Antarctic Peninsula., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Mercury (Hg) biomonitoring requires a precise understanding of the internal processes contributing to disparities between the Hg sources in the environment and the Hg measured in the biota. In this study, we investigated the use of Hg stable isotopes to trace Hg accumulation in Adélie and emperor penguin chicks from four breeding colonies in Antarctica. Interspecific variation of Δ 199 Hg in penguin chicks reflects the distinct foraging habitats and Hg exposures in adults. Chicks at breeding sites where adult penguins predominantly consumed mesopelagic prey showed relatively lower Δ 199 Hg values than chicks that were primarily fed epipelagic krill. Substantial δ 202 Hg variations in chick tissues were observed in both species (Adélie: -0.11 to 1.13‰, emperor: -0.27 to 1.15‰), whereas only emperor penguins exhibited the lowest δ 202 Hg in the liver and the highest in the feathers. Our results indicate that tissue-specific δ 202 Hg variations and their positive correlations with % MeHg resulted from MeHg demethylation in the liver and kidneys of emperor penguin chicks, whereas Adélie penguin chicks showed different internal responses depending on their exposure to dietary MeHg. This study highlights the importance of considering intra- and interspecific variations in adult foraging ecology and MeHg demethylation when selecting penguin chicks for Hg biomonitoring.

In vertebrates, fasting is an intricate physiological process associated with strong metabolic changes, yet its effect on pollutant residue variation is poorly understood. Here, we quantified long-term changes in plasma concentrations of 20 organochlorine and 16 perfluoroalkyl pollutants in king penguins Aptenodytes patagonicus during the breeding and molting fasts, which are marked by low and high levels of protein catabolism, respectively, and by strong lipid use. The profile of measured pollutants in plasma was dominated by perfluorooctanesulfonic acid (PFOS, initial relative contribution of 60%). Initial total pollutant concentrations were similar in molting (3.3-5.7 ng g -1 ww) and breeding penguins (range of 4.2-7.3 ng g -1 wet weight, ww). Long-term fasting (25 days) for molting and breeding led, respectively, to a 1.8- and 2.2-fold increase in total plasma pollutant concentrations, although the rate and direction of change were compound-specific. Hexachlorbenzene (HCB) and PFOS concentrations increased in plasma (net mobilization) during both types of fasting, likely due to lipid use. Plasma perfluoroundecanoate (PFUnDA) and perfluorotridecanoate (PFTrDA) concentrations increased in breeders (net mobilization) but decreased in molting individuals (net excretion), suggesting a significant incorporation of these pollutants into feathers. This study is a key contribution to our understanding of pollutant variation in blood during long-term fasting in wildlife.

Electrophoresis is a useful diagnostic tool for detecting inflammation, including inflammation associated with infectious diseases (eg, aspergillosis in penguins). To our knowledge, reference intervals are not available for plasma proteins via electrophoresis in Humboldt penguins ( Spheniscus humboldti ). Therefore, preliminary reference intervals for blood plasma proteins measured by capillary zone electrophoresis were calculated for Humboldt penguins from a single zoological collection, and possible differences between the sexes and the ages of the birds were evaluated. Lithium heparinized plasma samples from 39 Humboldt penguins were analyzed. The following sex- and age-independent reference intervals were calculated: total protein 33.8-70.4 g/L, prealbumin 1.9-4.9 g/L, albumin 12.9-31.1 g/L, albumin: globulin ratio 0.7-1.7, α-globulins 4.5-11.6 g/L, β-globulins 5.6-20.6 g/L, and γ-globulins 2.6-8.4 g/L. Male penguins had a significantly ( P = 0.047) higher albumin: globulin ratio and lower percentage of β-globulins ( P = 0.015) in comparison with female penguins. Prealbumin (g/L) significantly ( P = 0.021) decreased with increased age of the penguins. These results showed some differences between the sexes and ages of the penguins, which should be considered when interpreting the results. Further studies are needed to determine whether differences in other age groups or seasons exist, and also to evaluate which infectious diseases affect plasma proteins and how the reference values calculated here may deviate in ill penguins.

The egg production of captive African penguins differs considerably between individuals. An understanding of the physiological differences in African penguins with relatively greater and lesser egg production is meaningful for the captive breeding program of this endangered species. The objective of this study was to investigate differential microbial composition and metabolites in captive African penguins with different egg production. Fecal samples were collected from captive female African penguins during the breeding season. The results of 16 S rRNA gene sequencing showed that African penguins with different egg production had similar microbial diversities, whereas a significant difference was observed between their microbial community structure. African penguins with relatively greater egg production exhibited a higher relative abundance of Alphaproteobacteria, Rhizobiales, Bradyrhizobiaceae, Bradyrhizobium and Bosea. Meanwhile, penguins with relatively lesser egg production had an increased proportion of Klebsiella and Plesiomonas. We further identified a total of 1858 metabolites in female African penguins by liquid chromatography-mass spectrometry analysis. Among these metabolites, 13 kinds of metabolites were found to be significantly differential between African penguins with different egg production. In addition, the correlation analysis revealed that the egg production had significant correlations with most of the differential microbial bacteria and metabolites. Our findings might aid in understanding the potential mechanism underlying the phenomenon of abnormal egg production in captive African penguins, and provide novel insights into the relationship between gut microbiota and reproduction in penguins., Competing Interests: Declaration of Competing Interest The authors declare that they have no competing interests., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Antarctica is the most remote and coldest regions of the planet, but the presence of REEs there has received little attention. This study assessed REE-contents in the feathers of adult gentoo penguins from Ardley Island, Kopaitic Island and Base O'Higgins. Field work was accomplished during 2011 (austral summer), and determination of elements was performed with ICP-MS. In general, REE-levels showed descending relations as follows: Ce > La > Y > Nd > Sc > Pr > Gd > Sm > Dy >Er > Yb > Eu > Ho > Tb > Tm > Lu. The data showed an increase of the levels of REEs from the lower part of the feather to the tip. This finding seems to be spatially dependent, but geochemical, anthropogenic conditions, feeding habits, sex, or even health status of birds should also be considered. It is a subject that requires deeper attention in future studies., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Humboldt Penguin (Spheniscus humboldti) population declines are attributable to several multifaceted anthropogenic impacts. At present, the exposure of Humboldt Penguins to high concentrations of heavy metals in the marine environment is a preeminent concern, due to mining along the Peruvian coast near key rookery sites. Metal and selenium concentrations were determined in eggs collected from September 2020 to April 2021 from a managed-care penguin population at the Brookfield Zoo to establish reference values for health indices conducted on wild populations. Concentrations of 16 elements, with emphasis on those found in mine efflux-arsenic, cadmium, copper, lead, mercury, selenium, and zinc-were assessed via inductively coupled plasma mass spectrometry in yolk, albumen, and eggshell. Data analyses indicate a clear delineation between egg constituents, with lipid-rich yolk displaying notably higher concentrations (μg/g) of arsenic (0.20 ± 0.064), chromium (0.086 ± 0.03), cobalt (0.01 ± 0.003), iron (238.65 ± 54.72), lead (0.32 ± 0.97), manganese (2.71 ± 0.66), molybdenum (0.57 ± 0.14), tin (3.29 ± 0.99), and zinc (64.03 ± 13.01) than other components (albumen and eggshell). These data confirm that heavy metals are partitioned differently across Humboldt Penguin egg components, which provides insight into the potential connection between embryonic nutrient source contamination and subsequent chick viability., (© Wildlife Disease Association 2024.)