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Albatrosses are among the most intensely studied groups of living birds, yet their fossil record remains sparse. Despite modern albatrosses being more abundant and widespread in the Southern Hemisphere, the vast majority of fossil albatrosses identified to date come from Northern Hemisphere localities. Here, we describe Plotornis archaeonautes sp. nov., a new albatross species from the earliest Miocene that represents the earliest record of Procellariiformes in New Zealand and the earliest uncontroversial record of the clade Pan-Diomedeidae from the Southern Hemisphere. Phylogenetic analyses support the placement of Plotornis outside of the clade uniting all extant albatrosses. The new fossil reveals that stem lineage albatrosses were widespread by the onset of the Neogene. Although the humerus of Plotornis archaeonautes exhibits a short processus supracondylaris dorsalis, this early species may have possessed at least one of the unique ossifications associated with the patagial bracing system present in modern albatrosses., Competing Interests: No potential conflict of interest was reported by the authors., (© 2023 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.)

Penguins lost the ability to fly more than 60 million years ago, subsequently evolving a hyper-specialized marine body plan. Within the framework of a genome-scale, fossil-inclusive phylogeny, we identify key geological events that shaped penguin diversification and genomic signatures consistent with widespread refugia/recolonization during major climate oscillations. We further identify a suite of genes potentially underpinning adaptations related to thermoregulation, oxygenation, diving, vision, diet, immunity and body size, which might have facilitated their remarkable secondary transition to an aquatic ecology. Our analyses indicate that penguins and their sister group (Procellariiformes) have the lowest evolutionary rates yet detected in birds. Together, these findings help improve our understanding of how penguins have transitioned to the marine environment, successfully colonizing some of the most extreme environments on Earth., (© 2022. The Author(s).)

Relative brain size has long been considered a reflection of cognitive capacities and has played a fundamental role in developing core theories in the life sciences. Yet, the notion that relative brain size validly represents selection on brain size relies on the untested assumptions that brain-body allometry is restrained to a stable scaling relationship across species and that any deviation from this slope is due to selection on brain size. Using the largest fossil and extant dataset yet assembled, we find that shifts in allometric slope underpin major transitions in mammalian evolution and are often primarily characterized by marked changes in body size. Our results reveal that the largest-brained mammals achieved large relative brain sizes by highly divergent paths. These findings prompt a reevaluation of the traditional paradigm of relative brain size and open new opportunities to improve our understanding of the genetic and developmental mechanisms that influence brain size., (Copyright © 2021 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

Feathers are the most complex integumentary structures in the animal world. They come in a variety of forms, the most familiar of which are remiges (flight feathers). Flight feathers are composed of a central shaft made up of a hollow calamus (quill), which is inserted into the skin, and a more distal rachis. Hundreds of parallel barbs branch from the sides of the rachis. In turn, smaller hooked barbules branch off the barbs, allowing them to interlock in a tight zipper-like fashion to form vanes. Variations in rachis, barb and barbule morphology result in other feather types such as contour feathers, bristles and down feathers. Feathers have a remarkable array of functions - they form airfoils and elaborate display structures, they serve to camouflage and insulate, to generate and help detect sound, and even to disintegrate into powder to condition other feathers., (Copyright © 2020 Elsevier Inc. All rights reserved.)

New Zealand is a globally significant hotspot for seabird diversity, but the sparse fossil record for most seabird lineages has impeded our understanding of how and when this hotspot developed. Here, we describe multiple exceptionally well-preserved specimens of a new species of penguin from tightly dated (3.36-3.06 Ma) Pliocene deposits in New Zealand. Bayesian and parsimony analyses place Eudyptes atatu sp. nov. as the sister species to all extant and recently extinct members of the crested penguin genus Eudyptes . The new species has a markedly more slender upper beak and mandible compared with other Eudyptes penguins. Our combined evidence approach reveals that deep bills evolved in both crested and stiff-tailed penguins ( Pygoscelis ) during the Pliocene. That deep bills arose so late in the greater than 60 million year evolutionary history of penguins suggests that dietary shifts may have occurred as wind-driven Pliocene upwelling radically restructured southern ocean ecosystems. Ancestral area reconstructions using BioGeoBEARS identify New Zealand as the most likely ancestral area for total-group penguins, crown penguins and crested penguins. Our analyses provide a timeframe for recruitment of crown penguins into the New Zealand avifauna, indicating this process began in the late Neogene and was completed via multiple waves of colonizing lineages.

Relative brain sizes in birds can rival those of primates, but large-scale patterns and drivers of avian brain evolution remain elusive. Here, we explore the evolution of the fundamental brain-body scaling relationship across the origin and evolution of birds. Using a comprehensive dataset sampling> 2,000 modern birds, fossil birds, and theropod dinosaurs, we infer patterns of brain-body co-variation in deep time. Our study confirms that no significant increase in relative brain size accompanied the trend toward miniaturization or evolution of flight during the theropod-bird transition. Critically, however, theropods and basal birds show weaker integration between brain size and body size, allowing for rapid changes in the brain-body relationship that set the stage for dramatic shifts in early crown birds. We infer that major shifts occurred rapidly in the aftermath of the Cretaceous-Paleogene mass extinction within Neoaves, in which multiple clades achieved higher relative brain sizes because of a reduction in body size. Parrots and corvids achieved the largest brains observed in birds via markedly different patterns. Parrots primarily reduced their body size, whereas corvids increased body and brain size simultaneously (with rates of brain size evolution outpacing rates of body size evolution). Collectively, these patterns suggest that an early adaptive radiation in brain size laid the foundation for subsequent selection and stabilization., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Our understanding of the earliest stages of crown bird evolution is hindered by an exceedingly sparse avian fossil record from the Mesozoic era. The most ancient phylogenetic divergences among crown birds are known to have occurred in the Cretaceous period 1-3 , but stem-lineage representatives of the deepest subclades of crown birds-Palaeognathae (ostriches and kin), Galloanserae (landfowl and waterfowl) and Neoaves (all other extant birds)-are unknown from the Mesozoic era. As a result, key questions related to the ecology 4,5 , biogeography 3,6,7 and divergence times 1,8-10 of ancestral crown birds remain unanswered. Here we report a new Mesozoic fossil that occupies a position close to the last common ancestor of Galloanserae and fills a key phylogenetic gap in the early evolutionary history of crown birds 10,11 . Asteriornis maastrichtensis, gen. et sp. nov., from the Maastrichtian age of Belgium (66.8-66.7 million years ago), is represented by a nearly complete, three-dimensionally preserved skull and associated postcranial elements. The fossil represents one of the only well-supported crown birds from the Mesozoic era 12 , and is the first Mesozoic crown bird with well-represented cranial remains. Asteriornis maastrichtensis exhibits a previously undocumented combination of galliform (landfowl)-like and anseriform (waterfowl)-like features, and its presence alongside a previously reported Ichthyornis-like taxon from the same locality 13 provides direct evidence of the co-occurrence of crown birds and avialan stem birds. Its occurrence in the Northern Hemisphere challenges biogeographical hypotheses of a Gondwanan origin of crown birds 3 , and its relatively small size and possible littoral ecology may corroborate proposed ecological filters 4,5,9 that influenced the persistence of crown birds through the end-Cretaceous mass extinction.

Background: Penguins (Sphenisciformes) are a remarkable order of flightless wing-propelled diving seabirds distributed widely across the southern hemisphere. They share a volant common ancestor with Procellariiformes close to the Cretaceous-Paleogene boundary (66 million years ago) and subsequently lost the ability to fly but enhanced their diving capabilities. With ∼20 species among 6 genera, penguins range from the tropical Galápagos Islands to the oceanic temperate forests of New Zealand, the rocky coastlines of the sub-Antarctic islands, and the sea ice around Antarctica. To inhabit such diverse and extreme environments, penguins evolved many physiological and morphological adaptations. However, they are also highly sensitive to climate change. Therefore, penguins provide an exciting target system for understanding the evolutionary processes of speciation, adaptation, and demography. Genomic data are an emerging resource for addressing questions about such processes., Results: Here we present a novel dataset of 19 high-coverage genomes that, together with 2 previously published genomes, encompass all extant penguin species. We also present a well-supported phylogeny to clarify the relationships among penguins. In contrast to recent studies, our results demonstrate that the genus Aptenodytes is basal and sister to all other extant penguin genera, providing intriguing new insights into the adaptation of penguins to Antarctica. As such, our dataset provides a novel resource for understanding the evolutionary history of penguins as a clade, as well as the fine-scale relationships of individual penguin lineages. Against this background, we introduce a major consortium of international scientists dedicated to studying these genomes. Moreover, we highlight emerging issues regarding ensuring legal and respectful indigenous consultation, particularly for genomic data originating from New Zealand Taonga species., Conclusions: We believe that our dataset and project will be important for understanding evolution, increasing cultural heritage and guiding the conservation of this iconic southern hemisphere species assemblage., (© The Author(s) 2019. Published by Oxford University Press.)

Avian diversification has been influenced by global climate change, plate tectonic movements, and mass extinction events. However, the impact of these factors on the diversification of the hyperdiverse perching birds (passerines) is unclear because family level relationships are unresolved and the timing of splitting events among lineages is uncertain. We analyzed DNA data from 4,060 nuclear loci and 137 passerine families using concatenation and coalescent approaches to infer a comprehensive phylogenetic hypothesis that clarifies relationships among all passerine families. Then, we calibrated this phylogeny using 13 fossils to examine the effects of different events in Earth history on the timing and rate of passerine diversification. Our analyses reconcile passerine diversification with the fossil and geological records; suggest that passerines originated on the Australian landmass ∼47 Ma; and show that subsequent dispersal and diversification of passerines was affected by a number of climatological and geological events, such as Oligocene glaciation and inundation of the New Zealand landmass. Although passerine diversification rates fluctuated throughout the Cenozoic, we find no link between the rate of passerine diversification and Cenozoic global temperature, and our analyses show that the increases in passerine diversification rate we observe are disconnected from the colonization of new continents. Taken together, these results suggest more complex mechanisms than temperature change or ecological opportunity have controlled macroscale patterns of passerine speciation., Competing Interests: The authors declare no conflict of interest., (Copyright © 2019 the Author(s). Published by PNAS.)

The emergence of islands has been linked to spectacular radiations of diverse organisms. Although penguins spend much of their lives at sea, they rely on land for nesting, and a high proportion of extant species are endemic to geologically young islands. Islands may thus have been crucial to the evolutionary diversification of penguins. We test this hypothesis using a fossil-calibrated phylogeny of mitochondrial genomes (mitogenomes) from all extant and recently extinct penguin taxa. Our temporal analysis demonstrates that numerous recent island-endemic penguin taxa diverged following the formation of their islands during the Plio-Pleistocene, including the Galápagos (Galápagos Islands), northern rockhopper (Gough Island), erect-crested (Antipodes Islands), Snares crested (Snares) and royal (Macquarie Island) penguins. Our analysis also reveals two new recently extinct island-endemic penguin taxa from New Zealand's Chatham Islands: Eudyptes warhami sp. nov. and a dwarf subspecies of the yellow-eyed penguin, Megadyptes antipodes richdalei ssp. nov. Eudyptes warhami diverged from the Antipodes Islands erect-crested penguin between 1.1 and 2.5 Ma, shortly after the emergence of the Chatham Islands (∼3 Ma). This new finding of recently evolved taxa on this young archipelago provides further evidence that the radiation of penguins over the last 5 Ma has been linked to island emergence. Mitogenomic analyses of all penguin species, and the discovery of two new extinct penguin taxa, highlight the importance of island formation in the diversification of penguins, as well as the extent to which anthropogenic extinctions have affected island-endemic taxa across the Southern Hemisphere's isolated archipelagos., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Beak shape plays a key role in avian radiations and is one of the most intensely studied aspects of avian evolution and ecology [1-4]. Perhaps no other group is more closely associated with the study of beak shape than Passeriformes (passerines or perching birds), the most species-rich ordinal clade of modern birds. However, despite their extraordinary present-day diversity, our understanding of early passerine evolution has been hindered by their sparse fossil record [5, 6]. Here, we describe two new species of early Eocene stem passerines from the Green River Formation of the United States and the Messel Formation of Germany. These species are the oldest fossil birds to exhibit a finch-like beak and provide the earliest evidence for a diet focused on small, hard seeds in crown birds. Given that granivory is a key adaptation that allows passerines to exploit open temperate environments, it is notable that both species occurred in subtropical environments [7, 8]. Phylogenetic analyses place both species within the Psittacopedidae, an extinct Eocene clade of zygodactyl stem passeriforms that also includes the slender-beaked nectarivorous Pumiliornis, the short-beaked Psittacopes, and the thrush-beaked Morsoravis. Our results reveal that stem passerines attained a diversity of beak shapes paralleling many of the morphotypes present in extant passerine finches, thrushes, and sunbirds, more than 35 million years before these morphotypes arose in the crown group. Extinction of these ecologically diverse fossil taxa may be linked to more sophisticated nest construction in anisodactyl crown passerines versus cavity-nesting in Eocene zygodactyl stem passerines [9]., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Evidence is accumulating for a rapid diversification of birds following the K-Pg extinction. Recent molecular divergence dating studies suggest that birds radiated explosively during the first few million years of the Paleocene; however, fossils from this interval remain poorly represented, hindering our understanding of morphological and ecological specialization in early neoavian birds. Here we report a small fossil bird from the Nacimiento Formation of New Mexico, constrained to 62.221-62.517 Ma. This partial skeleton represents the oldest arboreal crown group bird known. Phylogenetic analyses recovered Tsidiiyazhi abini gen. et sp. nov. as a member of the Sandcoleidae, an extinct basal clade of stem mousebirds (Coliiformes). The discovery of Tsidiiyazhi pushes the minimum divergence ages of as many as nine additional major neoavian lineages into the earliest Paleocene, compressing the duration of the proposed explosive post-K-Pg radiation of modern birds into a very narrow temporal window parallel to that suggested for placental mammals. Simultaneously, Tsidiiyazhi provides evidence for the rapid morphological (and likely ecological) diversification of crown birds. Features of the foot indicate semizygodactyly (the ability to facultatively reverse the fourth pedal digit), and the arcuate arrangement of the pedal trochleae bears a striking resemblance to the conformation in owls (Strigiformes). Inclusion of fossil taxa and branch length estimates impacts ancestral state reconstructions, revealing support for the independent evolution of semizygodactyly in Coliiformes, Leptosomiformes, and Strigiformes, none of which is closely related to extant clades exhibiting full zygodactyly., Competing Interests: The authors declare no conflict of interest.

The rapidly expanding interest in, and availability of, digital tomography data to visualize casts of the vertebrate endocranial cavity housing the brain (endocasts) presents new opportunities and challenges to the field of comparative neuroanatomy. The opportunities are many, ranging from the relatively rapid acquisition of data to the unprecedented ability to integrate critically important fossil taxa. The challenges consist of navigating the logistical barriers that often separate a researcher from high-quality data and minimizing the amount of non-biological variation expressed in endocasts - variation that may confound meaningful and synthetic results. Our purpose here is to outline preferred approaches for acquiring digital tomographic data, converting those data to an endocast, and making those endocasts as meaningful as possible when considered in a comparative context. This review is intended to benefit those just getting started in the field but also serves to initiate further discussion between active endocast researchers regarding the best practices for advancing the discipline. Congruent with the theme of this volume, we draw our examples from birds and the highly encephalized non-avian dinosaurs that comprise closely related outgroups along their phylogenetic stem lineage., (© 2015 Anatomical Society.)

Substantial changes in bone histology accompany the secondary adaptation to life in the water. This transition is well documented in several lineages of mammals and non-avian reptiles, but has received relatively little attention in birds. This study presents new observations on the long bone microstructure of penguins, based on histological sections from two extant taxa (Spheniscus and Aptenodytes) and eight fossil specimens belonging to stem lineages (†Palaeospheniscus and several indeterminate Eocene taxa). High bone density in penguins results from compaction of the internal cortical tissues, and thus penguin bones are best considered osteosclerotic rather than pachyostotic. Although the oldest specimens sampled in this study represent stages of penguin evolution that occurred at least 25 million years after the loss of flight, major differences in humeral structure were observed between these Eocene stem taxa and extant taxa. This indicates that the modification of flipper bone microstructure continued long after the initial loss of flight in penguins. It is proposed that two key transitions occurred during the shift from the typical hollow avian humerus to the dense osteosclerotic humerus in penguins. First, a reduction of the medullary cavity occurred due to a decrease in the amount of perimedullary osteoclastic activity. Second, a more solid cortex was achieved by compaction. In extant penguins and †Palaeospheniscus, most of the inner cortex is formed by rapid osteogenesis, resulting an initial latticework of woven-fibered bone. Subsequently, open spaces are filled by slower, centripetal deposition of parallel-fibered bone. Eocene stem penguins formed the initial latticework, but the subsequent round of compaction was less complete, and thus open spaces remained in the adult bone. In contrast to the humerus, hindlimb bones from Eocene stem penguins had smaller medullary cavities and thus higher compactness values compared with extant taxa. Although cortical lines of arrested growth have been observed in extant penguins, none was observed in any of the current sampled specimens. Therefore, it is likely that even these 'giant' penguin taxa completed their growth cycle without a major pause in bone deposition, implying that they did not undergo a prolonged fasting interval before reaching adult size., (© 2015 Anatomical Society.)

Remarkable feathered dinosaur fossils have blurred the lines between early birds and their non-avian dinosaur relatives. Rapid skeletal evolution and decreasing body size along one particular lineage of theropod dinosaurs paved the way for the spectacular radiation of birds., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Major disparities are recognized between molecular divergence dates and fossil ages for critical nodes in the Tree of Life, but broad patterns and underlying drivers remain elusive. We harvested 458 molecular age estimates for the stem and crown divergences of 67 avian clades to explore empirical patterns between these alternate sources of temporal information. These divergence estimates were, on average, over twice the age of the oldest fossil in these clades. Mitochondrial studies yielded older ages than nuclear studies for the vast majority of clades. Unexpectedly, disparity between molecular estimates and the fossil record was higher for divergences within major clades (crown divergences) than divergences between major clades (stem divergences). Comparisons of dates from studies classed by analytical methods revealed few significant differences. Because true divergence ages can never be known with certainty, our study does not answer the question of whether fossil gaps or molecular dating error account for a greater proportion of observed disparity. However, empirical patterns observed here suggest systemic overestimates for shallow nodes in existing molecular divergence dates for birds. We discuss underlying biases that may drive these patterns., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Pelagornithidae is an extinct clade of birds characterized by bizarre tooth-like bony projections of the jaws. Here, the flight capabilities of pelagornithids are explored based on data from a species with the largest reported wingspan among birds. Pelagornis sandersi sp. nov. is represented by a skull and substantial postcranial material. Conservative wingspan estimates (∼6.4 m) exceed theoretical maximums based on extant soaring birds. Modeled flight properties indicate that lift:drag ratios and glide ratios for P. sandersi were near the upper limit observed in extant birds and suggest that pelagornithids were highly efficient gliders, exploiting a long-range soaring ecology.

A feathered specimen of a new species of Eocypselus from the Early Eocene Green River Formation of Wyoming provides insight into the wing morphology and ecology in an early part of the lineage leading to extant swifts and hummingbirds. Combined phylogenetic analysis of morphological and molecular data supports placement of Eocypselus outside the crown radiation of Apodiformes. The new specimen is the first described fossil of Pan-Apodiformes from the pre-Pleistocene of North America and the only reported stem taxon with informative feather preservation. Wing morphology of Eocypselus rowei sp. nov. is intermediate between the short wings of hummingbirds and the hyper-elongated wings of extant swifts, and shows neither modifications for the continuous gliding used by swifts nor modifications for the hovering flight style used by hummingbirds. Elongate hindlimb elements, particularly the pedal phalanges, also support stronger perching capabilities than are present in Apodiformes. The new species is the smallest bird yet described from the Green River Formation, and supports the hypothesis that a decrease in body size preceded flight specializations in Pan-Apodiformes. The specimen also provides the first instance of melanosome morphology preserved in association with skeletal remains from the Green River Formation.

Africa hosts a single breeding species of penguin today, yet the fossil record indicates that a diverse array of now-extinct taxa once inhabited southern African coastlines. Here, we show that the African penguin fauna had a complex history involving multiple dispersals and extinctions. Phylogenetic analyses and biogeographic reconstructions incorporating new fossil material indicate that, contrary to previous hypotheses, the four Early Pliocene African penguin species do not represent an endemic radiation or direct ancestors of the living Spheniscus demersus (blackfooted penguin). A minimum of three dispersals to Africa, probably assisted by the eastward-flowing Antarctic Circumpolar and South Atlantic currents, occurred during the Late Cenozoic. As regional sea-level fall eliminated islands and reduced offshore breeding areas during the Pliocene, all but one penguin lineage ended in extinction, resulting in today's depleted fauna.

Penguins (Sphenisciformes) inhabit some of the most extreme environments on Earth. The 60+ Myr fossil record of penguins spans an interval that witnessed dramatic shifts in Cenozoic ocean temperatures and currents, indicating a long interplay between penguin evolution and environmental change. Perhaps the most celebrated example is the successful Late Cenozoic invasion of glacial environments by crown clade penguins. A major adaptation that allows penguins to forage in cold water is the humeral arterial plexus, a vascular counter-current heat exchanger (CCHE) that limits heat loss through the flipper. Fossil evidence reveals that the humeral plexus arose at least 49 Ma during a 'Greenhouse Earth' interval. The evolution of the CCHE is therefore unrelated to global cooling or development of polar ice sheets, but probably represents an adaptation to foraging in subsurface waters at temperate latitudes. As global climate cooled, the CCHE was key to invasion of thermally more demanding environments associated with Antarctic ice sheets.

Background: The early Eocene Green River Formation avifauna preserves exceptional exemplars of the earliest unambiguous stem representatives of many extant avian clades. We identify the basal-most member of Podargiformes (extant and fossil stem lineage frogmouths) based on a new specimen of Fluvioviridavis platyrhamphus, a unique neoavian bird from the Fossil Butte Member of the Green River Formation of Wyoming. Extant frogmouths (Podargidae) comprise approximately 13 nocturnal species with an exclusively Australasian distribution., Methodology/principal Findings: The new specimen was included in a combined phylogenetic analysis of morphological (osteology and soft tissue) and molecular sequence (cytochrome b, c-myc exon 3, and RAG) data sampling species-level taxa from both extant and extinct members of Steatornithidae, Podargidae, Caprimulgidae, Nyctibiidae, Aegothelidae, and Apodiformes ( = Strisores). New data from F. platyrhamphus help resolve phylogenetic relationships within Strisores, supporting placement of F. platyrhamphus and the European fossil form Masillapodargus longipes as basal parts of Podargiformes and also supporting a sister taxon relationship between Podargiformes and Steatornithiformes (oilbirds) within Strisores. This relationship is recovered only when fossil taxa are included, reaffirming the potential impact of stem fossil taxa on inferences of phylogenetic relationships. The well-preserved mandible and palate of the new specimen demonstrate that many of the unique characteristics of the skull that characterize the crown frogmouth clade Podargidae arose early in the evolutionary history of the clade, over 50 million years ago. Comparisons with the new specimen also indicate that Eurofluvioviridavis and Fluvioviridavis are not closely related., Conclusions/significance: Together with the European fossil frogmouth Masillapodargus, Fluvioviridavis shows that Podargiformes had a much wider geographic distribution in the past, whereas extant species are restricted to Australasia. The Eocene record of Strisores from the Green River Formation and Messel Formation indicates most major subclade divergences had already occurred by the early-middle Eocene.

Penguin feathers are highly modified in form and function, but there have been no fossils to inform their evolution. A giant penguin with feathers was recovered from the late Eocene (~36 million years ago) of Peru. The fossil reveals that key feathering features, including undifferentiated primary wing feathers and broad body contour feather shafts, evolved early in the penguin lineage. Analyses of fossilized color-imparting melanosomes reveal that their dimensions were similar to those of non-penguin avian taxa and that the feathering may have been predominantly gray and reddish-brown. In contrast, the dark black-brown color of extant penguin feathers is generated by large, ellipsoidal melanosomes previously unknown for birds. The nanostructure of penguin feathers was thus modified after earlier macrostructural modifications of feather shape linked to aquatic flight.

Flexible, or soft-shelled, eggs are almost unknown in the fossil record, leaving large gaps in our knowledge of the reproductive biology of many tetrapod clades. Here, we report two flexible-shelled eggs of the hyphalosaurid choristodere Hyphalosaurus baitaigouensis from the Early Cretaceous of China, one containing an embryo and the second associated with a neonate. Choristoderes are an enigmatic group of aquatic reptiles that survived the K-T extinction but died out in the Miocene. Hyphalosaurids, a specialized clade of Choristodera, resemble miniature plesiosaurs and are considered to be primarily aquatic in habit. Scanning electron microscopy of samples from the eggs reveals a thin, non-columnar external mineralized layer characterized by rounded nodes and tentatively identified poorly structured irregular pores, with an underlying amorphous layer presumably representing decomposed protein fibrils. While the relationships of Choristodera remain controversial, eggshell microstructure more closely resembles that of Lepidosauromorpha (the lineage including lizards) as opposed to that of Archosauromorpha (the lineage including birds and crocodiles).

We present the first detailed description of the giant Eocene penguin Icadyptes salasi. The species is characterized by a narrow skull with a hyper-elongate spear-like beak, a robust cervical column and a powerful flipper. The bony beak tip of Icadyptes is formed by fusion of several elements and is unique among penguins, differing markedly from previously described giant penguin beaks. Vascular canal patterning similar to that of boobies, frigatebirds and albatrosses suggests I. salasi may have had a thin, sheet-like rhamphotheca unlike the thick rugose rhamphotheca of modern penguins. Together, these features suggest a novel ecology for I. salasi, most likely involving the capture of larger prey items via spearing. As the first described giant penguin specimen to preserve a complete wing skeleton, the I. salasi holotype yields significant insight into the shape, proportions and orientation of the wing in giant penguins. In articulation, the forelimb of I. salasi is straighter, permitting less manus and antibrachium flexion, than previous depictions of giant penguin wings. Cross-sections of the humerus and ulna reveal a level of osteosclerosis equalling or surpassing that of extant penguins. Based on ontogenetic data from extant penguins and the morphology of the carpometacarpus of I. salasi, we infer the retention of a free alular phalanx in basal penguins. Previously, the status of this element in penguins was disputed. Differences in the proportions of the manual phalanges contribute to a more abruptly tapering wingtip in I. salasi compared with crown penguins. Fossils from Peru, including the I. salasi holotype specimen, document that penguins expanded to nearly the whole of their extant latitudinal range early in their evolutionary history and during one of the warmest intervals in the Cenozoic.

Although the long-necked choristodere Hyphalosaurus is the most abundant tetrapod fossil in the renowned Yixian Formation fossil beds of Liaoning Province, China, the genus has only been briefly described from largely unprepared specimens. This paper provides a thorough osteological description of the type species Hyphalosaurus lingyuanensis and the con-generic species Hyphalosaurus baitaigouensis based on the study of fossils from several research institutions in China. The diagnoses for these two species are revised based on comparison of a large sample of specimens from the type area and horizon of each of the two species. The skull, better known in H. baitaigouensis, exhibits key choristodere synapomorphies including an elongate contact between the prefrontals and posteriorly expanded supratemporal fenestrae that strongly support the placement of the highly derived hyphalosaurids within Choristodera. Both species of Hyphalosaurus share a proportionally small head, an elongate neck, a relatively unspecialized appendicular skeleton and a long, dorsoventrally heightened tail. Soft tissue preservation in several specimens provides rare insight into the integument of an extinct group. The integument of Hyphalosaurus is made up of small polygonal scales with several parasagittal rows of large, keeled, ovoid scutes. These possibly ornamental scutes bear a strong resemblance to the rows of large scutes in the monjurosuchid choristodere Monjurosuchus splendens. Observations from a variety of growth stages reveal that significant ontogenetic change in the proportions of the body and limb bones occurred in both species of Hyphalosaurus.

New penguin fossils from the Eocene of Peru force a reevaluation of previous hypotheses regarding the causal role of climate change in penguin evolution. Repeatedly it has been proposed that penguins originated in high southern latitudes and arrived at equatorial regions relatively recently (e.g., 4-8 million years ago), well after the onset of latest Eocene/Oligocene global cooling and increases in polar ice volume. By contrast, new discoveries from the middle and late Eocene of Peru reveal that penguins invaded low latitudes >30 million years earlier than prior data suggested, during one of the warmest intervals of the Cenozoic. A diverse fauna includes two new species, here reported from two of the best exemplars of Paleogene penguins yet recovered. The most comprehensive phylogenetic analysis of Sphenisciformes to date, combining morphological and molecular data, places the new species outside the extant penguin radiation (crown clade: Spheniscidae) and supports two separate dispersals to equatorial (paleolatitude approximately 14 degrees S) regions during greenhouse earth conditions. One new species, Perudyptes devriesi, is among the deepest divergences within Sphenisciformes. The second, Icadyptes salasi, is the most complete giant (>1.5 m standing height) penguin yet described. Both species provide critical information on early penguin cranial osteology, trends in penguin body size, and the evolution of the penguin flipper.