Thesis (Ph. D.)--University of Texas at Austin, 2007. / Vita. Includes bibliographical references.

The excellent penguin fossil record is temporally long, among the oldest of modern birds, and contains fossils on every Gondwanan continent except India. However, most of fifty-nine named taxa are isolated skeletal elements, many of which are noncomparable. Fossil diversity is highest in New Zealand, with additional Antarctic and Patagonian faunas and fewer remains from Australia and South Africa. Phylogenetic hypotheses place penguins within Aves and Neornithes, but further relationships remain contentious. Recent work clarified living species' phylogeny, but none examined all fossil taxa. I describe penguin skeletal anatomy using CT scans and museum specimens, providing the first such description for all living and extinct species in an explicitly phylogenetic framework. All elements are phylogenetically variable, intergeneric variation is large, and extinct taxa are more variable than extant. I recommend that future systematic works include all elements for extinct species diagnoses, osteology for living species, and discussions of intraspecific variation. This description grounds my phylogenetic analysis, based on a 503-character matrix of osteological, myological, integumentary, and behavioral characters. This greatly expands previous datasets, and allows recovery of a highly resolved phylogeny, including monophyly of two extinct clades and the crown-group. Data partitions support different levels of relationship, whereas missing data and outgroup choice drastically affect recovered topology. Incorporating the maximum amount of data gives the highest resolution by recovering all relevant character states. I propose the first formal phylogenetic nomenclature for sphenisciforms, and define and diagnose previously used terms such as Panspheniscidae (total group), Sphenisciformes (known penguins), Spheniscidae (crown-group), Palaeeudyptidae and Paraptenodytidae (two extinct clades). I coin Spheniscoidea (Spheniscidae + Paraptenodytidae) and Australodyptinae (Aptenodytes + Pygoscelis). After calibration with geologic time, I calculate the completeness of the pencuin record and determine confidence intervals to estimate a Cretaceous origin and an Eocene crown-clade origin. These dates and the derived phylogenetic placement of penguins suggest that numerous extant bird lineages may also have Cretaceous origins. I recover a New Zealand origin for penguins and a West Antarctic origin for the crown-group and extinct clades. Correlating penguins with tectonic and climatic data indicates an Eocene adaptive radiation, probably related to latest-stage Gondwanan breakup and associated global cooling.

Northern Rockhopper penguin populations, Eudyptes moseleyi, are declining globally, and at Tristan da Cunha have undergone severe declines (> 90% in the last 130 years), the cause(s)of which are unknown. There is a paucity of data on this species in the South Atlantic Ocean, therefore their trophic ecology at Tristan da Cunha was studied, specifically focusing on diet, using stomach content analysis and stable isotope analysis (SIA), in conjunction with an analysis of diving behaviour, assessed using temperature-depth recorders. In order to evaluate the influence of gender on foraging, a morphometric investigation of sexual dimorphism was confirmed using molecular analysis. Additionally, plasma corticosterone levels were measured to examine breeding stage and presence of blood parasites as potential sources of stress during the breeding season. Northern Rockhopper penguins at Tristan da Cunha displayed a high degree of foraging plasticity, and fed opportunistically on a wide variety of prey, probably reflecting local small-scale changes in prey distribution. Zooplankton dominated (by mass) the diet of guard stage females, whereas small meso-pelagic fish (predominantly Photichthyidae)dominated diet of adults of both sexes in the crèche stage, with cephalopods contributing equally in both stages. Adults consistently fed chicks on lower-trophic level prey (assessed using SIA), probably zooplankton, than they consumed themselves indicating that the increasing demands of growing chicks were not met by adults through provisioning of higher- quality prey. SIA also indicated that adults foraged in different oceanic water masses when feeding for self-maintenance and for chick provisioning, thus temporally segregating the prey consumed for different purposes. It is possible that adults ‘selected’ these higher quality prey for themselves, or this may be a reflection of opportunistic behaviour. At Tristan da Cunha sexual dimorphism was observed in culmen dimensions (length, depth, width), with males having larger beaks and feeding on larger individuals of squid and fish than females. No sexual segregation in terms of foraging habitat (i.e. different water masses, based on ð¹³N or trophic level ð¹⁵N) during the breeding season or pre-moult period was revealed through SIA, and stomach content analysis revealed no sexual differences in prey species targeted. The results of SIA of feathers indicate that during the pre-moult period birds foraged in different water masses than during the breeding period. The fact that throughout the breeding season birds foraged in similar habitats suggests no intra-specific competition, despite both sexes feeding on the same prey.Birds were generally diurnal, daily foragers (12 – 16 hr trips), with extended trips (maximum duration 35.5 hours) and nocturnal diving recorded in a few individuals. Birds dived well within their physiological limits, predominantly utilising the upper 20m of the water column, employing two different strategies to target different prey items. Long, deep (30 – 40 m), energetically costly dives were performed when targeting energy-rich prey (fish), and a greater number of shorter, shallower (5 – 20 m), energy-efficient dives were performed when targeting prey with a lower energy content (zooplankton). More than half of the sampled study population were infected with the intra-cellular blood parasite Babesia, but infection showed no relationship to body mass, corticosterone levels or breeding success. Fasting birds showed no signs of elevated corticosterone levels, suggesting they had acquired sufficient fat reserves prior to breeding. Failed breeders did not exhibit elevated corticosterone levels. Tristan skuas, Catharacta antarctica hamiltoni, were observed to be a significant cause of egg and chick mortality. The absence of sex-based differences in foraging, and the absence of any signs of stress in relation to body mass, presence of Babesia or breeding stage, suggest that there are no obvious signs of high levels of stress or food limitations during breeding at Tristan da Cunha.

It is important to the survival of the Africa Penguin (Spheniscus demersus) population that breeding at the nest site is successful and that large numbers of chicks are fledged into the breeding population. Nest distribution on Bird Island is not random and locality preferences for breeding exist. Although it seems that sufficient area exists on Bird Island for penguin nests, it can hardly be considered as suitable to optimise breeding. During prolonged heat conditions, breeders relocated to nest sites that were sheltered. Nests that were below ground in burrows was the only habitat that did not suffer nest desertion whilst all the other habitat types (including those that were sheltered) experienced 2-3 fold declines in nest numbers. Nests density and the selection of suitable nest sites are significantly influenced by the stage of breeding that the majority of birds are in, yet nests that are shaded, well-ventilated and protected seem to be the most preferred sites for breeding. Adults that attempt to breed are considered then to be in a healthy condition and will usually lay a double clutch (Randell 1983). The frequency of double clutches being laid during the peak breeding attempt was significantly higher compared to the replacement one. Breeding failure was fairly similar to breeding success during the incubation stage for nests with double clutches however, was substantially higher in single clutches. The growth rate of chicks was best fit to the von Bertalanffy growth curve in 90 percent of the cases. The overall growth rate of chicks from double broods was faster than from single broods, however was not significant. A-chicks maitain a high growth rate until they fledged. Yet, the sibling B-chick recorded the lowest growth rate of the successfully fledged chicks and up to until day 30 recorded a similar weight to those chicks that failed to fledge. Contrary to findings of Randall (1983), chicks from single broods delayed fledging, recorded lowest overall growth rates and experienced the greatest weight loss of all groups, yet fledged successfully. In order for chicks to fledge successfully, they needed to obtain a weight of 1060 g before day 30.5 in their growth cycles to avoid death due to startvation later on. Single chicks that are raised from a double cluth, fledged more other than chicks raised from a single clutch. Unfit or ill-adapted breeders that are marginal in the capabilities of raising offspring, already manifest in a small clutch size and offspring unable to obtain adequate weights during the initial stages of growth.

No description available.

Penguins are middle- to large-sized sea birds and are widely distributed in the Southern Hemisphere. They have completely lost the capability for the aerial flight, but are highly efficient wing-propelled swimmers and divers. They have a long fossil record over 60 million years, and their origin could possibly extend back to the late Cretaceous. This study aims to elaborate the course of penguin evolution and driving force of changes based on fossil records of penguins. Numerous fossil penguin specimens have been collected and studied from New Zealand, Antarctica, South America, Australia, and Africa. Studies on fossil penguins have spanned about 150 years history since Huxley (1859). Previous works on fossil penguins have achieved excellent results, but at the same time, left considerable confusion on taxonomy and anatomical interpretation, mainly because of the poor nature of the penguin fossils in early studies. Examination of newly found materials and updated evaluation of previously studied materials are needed, using modern methods. During about 150 years of fossil penguins study since Huxley1859, more than 40 genera and 70 species have been described. The number of specimens listed in the published literature amounts to more than 1300. Chapter II reviews all those fossil penguins in a summarised and consistent style, aiming to present the taxonomy used in this study as a primary and essential resource for research. The chapter also provides other information on fossil penguins, such as geological data and an assessment of the skeletal association of the specimens referred to a species. Chapter III introduces the osteology of penguins, by describing and comparing the skeletal characteristics and variation of both extant and fossil species. Though previous works on penguins osteology are extensive, the interpretation of the homology, and resulted terminology, are occasionally inappropriate, or incorrect, because of the highly-specialised structure. Many of the new, yet undescribed, fossils prompt a comprehensive update of those previous studies, to understand the nature of morphological variation in penguins, and to correct or clarify confusion in previous works. The New Zealand fossil penguin fauna is one of the most significant for fossil penguin studies, but there are many undescribed fossil penguin specimens. Chapter IV provides accounts of such materials. Chapter IV also reviews previously-described New Zealand fossil penguins, usually re-evaluated using new materials. This chapter includes reassessment of the controversial, first-described fossil penguin Palaeeudyptes antarcticus, description of an enigmatic new species (Pakudyptes hakataramea gen. et sp. nov.) which could elucidate the evolutionary pattern of the penguin wing, description of new materials of Platydyptes revealing a unique structure and functional interpretation, and redescriptions with functional interpretation of Pachydyptes and Archaeospheniscus. Published relationships within penguins have not been adequately discussed but stated within rather rough frameworks, so that the relationships within penguins were unclear. Chapter V provides an explicit framework for the phylogeny of penguins. Osteology-based cladistic analysis was performed to seek out the relationships within penguins, using observations on both extant and fossil penguins. There are several important grades in penguin history, which are structurally distant from each other. Results also agree with the published views in which the extant penguins form a rigid group, but Simpson�s subfamily groupings are only partly supported. A postulated phylogenetic tree includes all known fossil penguin taxa including un-named ones. Chapter VI, as a synthesis of contents of previous chapters, provides a broad interpretation of penguin evolution through the Cenozoic: origin, body size increase, demise of 'giant penguins', and the emergence of modern penguins. The chapter gives a global picture of the interaction of penguins, pinnipeds, cetaceans, and temperature and sea-level change. Two main sections are: 'Origin of penguins' and 'Evolutionary process of penguins.' The loss of aerial flight and increase of body size were possibly triggered by the K/T mass extinction event which drastically reduced the predatory pressure for early penguins. The 'giant penguins' survived until the Late Oligocene but declined as the oceans modernised, and new forms of whales with advanced feeding function appeared. There is controversy about appearance of modern penguins. The fossil-based hypothesis (relatively recent origin for crown-penguins) contradicts the molecular-based one (ancient origin for crown penguins), though 'hard evidence' at present does not easily refute either hypothesis.

At the outbreak of the First World War, the archaic principles of nationalism and masculinity ruled Britain. These principles placed on men expectations that had become unrealistic due to the changed nature of warfare. The new horrors of war and the loss of the masculine characteristic of self-control produced a high frequency of combat trauma. For such victims of the war, the healing of psychological conditions required the assignment of meaning to their trauma, accomplished through the communication of loss to the civilian population. The problem was the inability of most non-combatants, including medical doctors, to comprehend ideas outside of the language-supported ideology that governed perception of reality. Instead of empathy, traumatized veterans were met with demands of conformity to the standards of masculinity established long before the war. Veterans who dissented from the official line of God, King and Country were silenced by the very society they fought to protect. Women writers, however, were free from the strictures of masculinity and were thus able to act as proxies to their counterparts. Rebecca West, Rose Macaulay, and Virginia Woolf challenged the dominant assumptions of war trauma and masculinity, each identifying language and anachronous ideology as the primary means used to promote conventional thought and silence discordance in society.

At the outbreak of the First World War, the archaic principles of nationalism and masculinity ruled Britain. These principles placed on men expectations that had become unrealistic due to the changed nature of warfare. The new horrors of war and the loss of the masculine characteristic of self-control produced a high frequency of combat trauma. For such victims of the war, the healing of psychological conditions required the assignment of meaning to their trauma, accomplished through the communication of loss to the civilian population. The problem was the inability of most non-combatants, including medical doctors, to comprehend ideas outside of the language-supported ideology that governed perception of reality. Instead of empathy, traumatized veterans were met with demands of conformity to the standards of masculinity established long before the war. Veterans who dissented from the official line of God, King and Country were silenced by the very society they fought to protect. Women writers, however, were free from the strictures of masculinity and were thus able to act as proxies to their counterparts. Rebecca West, Rose Macaulay, and Virginia Woolf challenged the dominant assumptions of war trauma and masculinity, each identifying language and anachronous ideology as the primary means used to promote conventional thought and silence discordance in society.

Penguins are middle- to large-sized sea birds and are widely distributed in the Southern Hemisphere. They have completely lost the capability for the aerial flight, but are highly efficient wing-propelled swimmers and divers. They have a long fossil record over 60 million years, and their origin could possibly extend back to the late Cretaceous. This study aims to elaborate the course of penguin evolution and driving force of changes based on fossil records of penguins. Numerous fossil penguin specimens have been collected and studied from New Zealand, Antarctica, South America, Australia, and Africa. Studies on fossil penguins have spanned about 150 years history since Huxley (1859). Previous works on fossil penguins have achieved excellent results, but at the same time, left considerable confusion on taxonomy and anatomical interpretation, mainly because of the poor nature of the penguin fossils in early studies. Examination of newly found materials and updated evaluation of previously studied materials are needed, using modern methods. During about 150 years of fossil penguins study since Huxley1859, more than 40 genera and 70 species have been described. The number of specimens listed in the published literature amounts to more than 1300. Chapter II reviews all those fossil penguins in a summarised and consistent style, aiming to present the taxonomy used in this study as a primary and essential resource for research. The chapter also provides other information on fossil penguins, such as geological data and an assessment of the skeletal association of the specimens referred to a species. Chapter III introduces the osteology of penguins, by describing and comparing the skeletal characteristics and variation of both extant and fossil species. Though previous works on penguins osteology are extensive, the interpretation of the homology, and resulted terminology, are occasionally inappropriate, or incorrect, because of the highly-specialised structure. Many of the new, yet undescribed, fossils prompt a comprehensive update of those previous studies, to understand the nature of morphological variation in penguins, and to correct or clarify confusion in previous works. The New Zealand fossil penguin fauna is one of the most significant for fossil penguin studies, but there are many undescribed fossil penguin specimens. Chapter IV provides accounts of such materials. Chapter IV also reviews previously-described New Zealand fossil penguins, usually re-evaluated using new materials. This chapter includes reassessment of the controversial, first-described fossil penguin Palaeeudyptes antarcticus, description of an enigmatic new species (Pakudyptes hakataramea gen. et sp. nov.) which could elucidate the evolutionary pattern of the penguin wing, description of new materials of Platydyptes revealing a unique structure and functional interpretation, and redescriptions with functional interpretation of Pachydyptes and Archaeospheniscus. Published relationships within penguins have not been adequately discussed but stated within rather rough frameworks, so that the relationships within penguins were unclear. Chapter V provides an explicit framework for the phylogeny of penguins. Osteology-based cladistic analysis was performed to seek out the relationships within penguins, using observations on both extant and fossil penguins. There are several important grades in penguin history, which are structurally distant from each other. Results also agree with the published views in which the extant penguins form a rigid group, but Simpson�s subfamily groupings are only partly supported. A postulated phylogenetic tree includes all known fossil penguin taxa including un-named ones. Chapter VI, as a synthesis of contents of previous chapters, provides a broad interpretation of penguin evolution through the Cenozoic: origin, body size increase, demise of 'giant penguins', and the emergence of modern penguins. The chapter gives a global picture of the interaction of penguins, pinnipeds, cetaceans, and temperature and sea-level change. Two main sections are: 'Origin of penguins' and 'Evolutionary process of penguins.' The loss of aerial flight and increase of body size were possibly triggered by the K/T mass extinction event which drastically reduced the predatory pressure for early penguins. The 'giant penguins' survived until the Late Oligocene but declined as the oceans modernised, and new forms of whales with advanced feeding function appeared. There is controversy about appearance of modern penguins. The fossil-based hypothesis (relatively recent origin for crown-penguins) contradicts the molecular-based one (ancient origin for crown penguins), though 'hard evidence' at present does not easily refute either hypothesis.

Thesis (M.I.N.C.) -- Lincoln University, 2009. / Also available via the World Wide Web.