Ecomorphological variation of the penguin wing.

Penguins (Aves, Sphenisciformes) are pursuit divers that feed mainly on krill, fish, and squid. Although they are opportunistic feeders, some species are more generalists than others and many show dietary preferences toward krill and other crustaceans or fish and squid. Their diving depth seems to follow a body size pattern and relates to the type of item that they prey on. Penguins dive with their wing; hence their wing musculature is responsible for the animal maneuverability and strength while diving. In the present study, ecological traits such as diving depths and prey composition are used to explore if morphology relates to foraging habits. A geometric morphometric approach is used to quantitatively address these morphological differences in the wing apparatus of all extant penguins and a fossil species taking into consideration allometric and phylogenetic factors. Results show that morphological differences among penguins with different diets are significant and strong; groups are well separated with the greatest differences found between piscivorous and crustacivorous penguins. Dive depth has a moderate covariation with morphology and a strong correspondence with wing area. Last, Madrynornis mirandus, an exceptionally well-preserved fossil from the Miocene of Patagonia, is found to be close to the piscivorous and generalist piscivorous species. It is proposed that swimming styles correlate with specific traits of the anatomy of wing and pectoral girdle skeleton and muscles.

Micro-CT guided illustration of the head anatomy of penguins (Aves: Sphenisciformes: Spheniscidae).

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.

Functional morphology of the cornea of the Little Penguin Eudyptula minor (Aves).

The cornea is a specialized component of the vertebrate eye that provides protection, refractive power, transparency for optical imaging and mechanical support. However, the corneas of birds have received little attention with no comprehensive study of their functional morphology. Using light microscopy and both scanning and transmission electron microscopy, the first description of the ultrastructure of all of the main components of the cornea in two different-sized individuals of the Little Penguin Eudyptula minor is presented. Two types of microprojections protrude from the surface of the cornea with a predominance of microridges and microvilli found in central (flattened) and peripheral regions, respectively. Epithelial cell density is higher in peripheral cornea, especially in the larger (older) individual, while there is a reduction of epithelial cell density with age. The cornea comprises a thick epithelium uniquely attached to the basement membrane with numerous incursions rather than anchoring fibres and anchoring plaques as is found in other vertebrate corneas. Posterior to Bowman's layer, the orthogonally-arranged collagen fibril lamellae in the stroma form extensive branches and anastomoses. Desçemet's membrane is well-developed with an anterior or foetal portion with long banding. However, the thickness of Desçemet's membrane is larger in the older individual with the inclusion of an additional irregular pale-staining posterior portion. Polygonal endothelial cells extend across the cornea as a monolayer with often tortuous cell junctions. Endothelial cell density increases towards the periphery, but decreases with age. Primary cilia are observed protruding through the central region of some endothelial cells into the anterior segment but subsurface structures resembling cilia suggest that these features may be more common. The ultrastructure of the corneal components reveals a range of functional adaptations that reflect the amphibious lifestyle of this seabird.

Existence and features of the myodural bridge in Gentoo penguins: A morphological study.

Recent studies have evidenced that the anatomical structure now known as the myodural bridge (MDB) connects the suboccipital musculature to the cervical spinal dura mater (SDM). In humans, the MDB passes through both the posterior atlanto-occipital and the posterior atlanto-axial interspaces. The existence of the MDB in various mammals, including flying birds (Rock pigeons and Gallus domesticus) has been previously validated. Gentoo penguins are marine birds, able to make 450 dives per day, reaching depths of up to 660 feet. While foraging, this penguin is able to reach speeds of up to 22 miles per hour. Gentoo penguins are also the world's fastest diving birds. The present study was therefore carried out to investigate the existence and characteristics of the MDB in Gentoo penguin (Pygoscelis papua), a non-flying, marine bird that can dive. For this study, six Gentoo penguin specimens were dissected to observe the existence and composition of their MDB. Histological staining was also performed to analyze the anatomic relationships and characteristic of the MDB in the Gentoo penguin. In this study, it was found that the suboccipital musculature in the Gentoo penguin consists of the rectus capitis dorsalis minor (RCDmi) muscle and rectus capitis dorsalis major (RCDma) muscle. Dense connective tissue fibers were observed connecting these two suboccipital muscles to the spinal dura mater (SDM). This dense connective tissue bridge consists of primarily type I collagen fibers. Thus, this penguin's MDB appears to be analogous to the MDB previously observed in humans. The present study evidences that the MDB not only exists in penguins but it also has unique features that distinguishes it from that of flying birds. Thus, this study advances the understanding of the morphological characteristics of the MDB in flightless, marine birds.

The skull of the Miocene Spheniscus urbinai (Aves, Sphenisciformes): osteology, brain morphology, and the cranial pneumatic systems.

Spheniscus urbinai represents one of four extinct Spheniscus species from the Cenozoic of southern South America, known from several poorly described diversely complete skulls and postcranial elements. Here, we present a review of the cranial osteology of all known specimens (collected in Argentina, Chile, and Peru), including a paleoneurological analysis using CT scans, and an exploration of its cranial pneumaticity compared to other extinct and living seabirds. Our results show that among Spheniscus species, S. urbinai exhibits slightly greater cranial pneumaticity than the living species. Additionally, we confirm previous findings which indicate that the marked reduction of cranial pneumaticity-which is characteristic of living penguins-occurred early during the Eocene (as observed in the Antarctic penguin MLP 12-I-20-1, but not in the coeval Anthropornis).

THE RADIOGRAPHIC AND ENDOSCOPIC ANATOMY AND DIGESTIVE MECHANISMS OF CAPTIVE AFRICAN PENGUINS (SPHENISCUS DEMERSUS).

The anatomy of the avian gastrointestinal (GI) tract is uniquely suited to each species' dietary requirements. African penguins (Spheniscus demersus) are charismatic and popular exhibit animals. As their prevalence grows, there is a need to understand their unique digestive tract to diagnose abnormalities. Reference material specific to the digestive tract of piscivores is scant, and knowledge of the GI tract of a healthy penguin is based on information from other birds. The purpose of this study is to determine the normal gross anatomy, transit time, and histopathologic structures of the penguin GI tract. Twelve clinically healthy penguins were selected for this study from the colony at the Maryland Zoo in Baltimore, which, at the time of this study, consisted of 55 birds. All penguins underwent a barium contrast study, and radiographic images were obtained until the entire GI tract was empty. Approximately 2 wk later, each penguin was anesthetized, and an endoscopic evaluation of the anterior GI tract was performed. Time from barium administration to defecation ranged from 17 to 70 min, and on average, barium clearance was 17.6 hr (range, 5-36 hr). Fluid from the ventriculus had an average pH of 2.75 and contained a mixed bacterial population. Koilin presence and thickness appreciated on endoscopy did not correspond with the thickness determined on histopathology. The results of this study provide a comparative baseline to use during diagnostic workups and help guide treatment decisions.

Symmetry breaking in the embryonic skin triggers directional and sequential plumage patterning.

The development of an organism involves the formation of patterns from initially homogeneous surfaces in a reproducible manner. Simulations of various theoretical models recapitulate final states of natural patterns, yet drawing testable hypotheses from those often remains difficult. Consequently, little is known about pattern-forming events. Here, we surveyed plumage patterns and their emergence in Galliformes, ratites, passerines, and penguins, together representing the three major taxa of the avian phylogeny, and built a unified model that not only reproduces final patterns but also intrinsically generates shared and varying directionality, sequence, and duration of patterning. We used in vivo and ex vivo experiments to test its parameter-based predictions. We showed that directional and sequential pattern progression depends on a species-specific prepattern: an initial break in surface symmetry launches a travelling front of sharply defined, oriented domains with self-organising capacity. This front propagates through the timely transfer of increased cell density mediated by cell proliferation, which controls overall patterning duration. These results show that universal mechanisms combining prepatterning and self-organisation govern the timely emergence of the plumage pattern in birds.

Flipper bone distribution reveals flexible trailing edge in underwater flying marine tetrapods.

Hydrofoil-shaped limbs (flipper-hydrofoils) have evolved independently several times in secondarily marine tetrapods and generally fall into two functional categories: (1) those that produce the majority of thrust during locomotion (propulsive flipper-hydrofoils); (2) those used primarily to steer and resist destabilizing movements such as yaw, pitch, and roll (controller flipper-hydrofoils). The morphological differences between these two types have been poorly understood. Theoretical and experimental studies on engineered hydrofoils suggest that flapping hydrofoils with a flexible trailing edge are more efficient at producing thrust whereas hydrofoils used in steering and stabilization benefit from a more rigid one. To investigate whether the trailing edge is generally more flexible in propulsive flipper-hydrofoils, we compared the bone distribution along the chord in both flipper types. The propulsive flipper-hydrofoil group consists of the forelimbs of Chelonioidea, Spheniscidae, and Otariidae. The controller flipper-hydrofoil group consists of the forelimbs of Cetacea. We quantified bone distribution from radiographs of species representing more than 50% of all extant genera for each clade. Our results show that the proportion of bone in both groups is similar along the leading edge (0-40% of the chord) but is significantly less along the trailing edge for propulsive flipper-hydrofoils (40-80% of the chord). Both flipper-hydrofoil types have little to no bony tissue along the very edge of the trailing edge (80-100% of the chord). This suggests a relatively flexible trailing edge for propulsive flipper-hydrofoils compared to controller flipper-hydrofoils in line with findings from prior studies. This study presents a morphological correlate for inferring flipper-hydrofoil function in extinct taxa and highlights the importance of a flexible trailing edge in the evolution of propulsive flipper-hydrofoils in marine tetrapods.

RADIOGRAPHIC EVALUATION OF CARDIAC SILHOUETTE IN CLINICALLY HEALTHY HUMBOLDT PENGUINS ( SPHENISCUS HUMBOLDTI).

Wild populations of Humboldt penguins ( Spheniscus humboldti) on the coasts of Chile and Peru have been declining because of food scarcity caused by the El Niño Southern Oscillation and human interference. Part of conserving this vulnerable and threatened species is maintaining the health of penguins within zoo collections. A variety of cardiovascular diseases has been reported in individuals from the Spheniscidae family including ventricular septal defects, Dirofilaria immitis infection, pulmonary hypertension, and valvular dysplasia, ultimately resulting in congestive heart failure. An accurate clinical picture of cardiovascular disease in Humboldt penguins requires diagnostics tailored to this specific species. The aim of this study was to establish a routine methodology for evaluating the cardiac silhouette of clinically healthy Humboldt penguins using vertebral heart scale (VHS), cardiocoelomic width ratio (CCWR), and a novel cardiac silhouette-to-keel ratio (CKR). Ventrodorsal and right lateral radiographs were taken of 10 mature Humboldt penguins during routine health evaluations. An echocardiographic exam of each penguin was performed to confirm that there was no evidence of cardiac structural remodeling from disease. Two penguins were excluded on the basis of echocardiographic findings; therefore, data from eight penguins were used to calculate objective cardiac measurements for the clinically healthy population. Right lateral radiographs were used to determine VHS (7.4-10.4) and CKR (3.4-4.4). Ventrodorsal radiographs were used for calculating CCWR (0.45-0.59). The excluded penguins had CCWRs that were outside the 95% confidence interval for the range generated by this study. This initial work supports that standardizing radiographic views provides objective measures for cardiac silhouette evaluation in this species. Further research in larger populations and comparison with birds having confirmed cardiac disease is needed to determine the value of these three measurement techniques in Humboldt penguins.

Morfologia do coração e dos vasos da base do pinguim-de-magalhães (Spheniscus magellanicus) / Morphology of heart and vessels of the cardiac base in Magellanic penguin (Spheniscus magellanicus)

O pinguim-de-magalhães é uma ave marinha de porte médio, de origem do hemisfério sul, com grandes colônias próximas à Patagônia. Em certas épocas do ano, alguns exemplares aparecem no litoral brasileiro, devido ao desvio de rotas de caça, e alguns indivíduos não conseguem retornar por debilidades na saúde. Foram utilizados 34 exemplares de Spheniscus magellanicus que vieram a óbito no litoral do estado de Espírito Santo. O presente estudo analisou a morfologia de câmaras e paredes cardíacas, valvas e artérias da base. Os fragmentos dessas regiões foram analisados histologicamente com coloração hematoxilina-eosina (HE) e Tricrômico de Gomori (TG), além da coloração Picrosirius Red (PSR) sob luz polarizada, visando observar, principalmente, a composição do tipo de colágeno existente em cada região descrita. Entre os 34 exemplares, nenhum apresentou discrepância em relação a sua morfologia. A tipificação do colágeno dessas regiões pelas colorações TG e PSR sob luz polarizada demonstrou a presença do colágeno tipo I em maior evidência que o tipo III, encontrada na maioria das estruturas, o que atribuiu a aparência avermelhada intensa a quase todas elas. Pode-se concluir que a anatomia cardíaca do pinguim-de-magalhães é semelhante à de outras aves, com predominância do colágeno do tipo I.(AU)

Magellanic penguin is medium-sized seabird originated from southern hemisphere with colonies near Patagonia. At certain times of the year in Brazilian coast, a few penguins lose their hunting routes and can´t return because they are very sick. Thirty-four penguins died in Espírito Santo´s coast. This study analyzed the cardiac morphology and morph metric of heart chambers and walls, valves, and arteries of the cardiac base. These parts were analysed and stained by Hematoxilin and eosin and Gomori´s trichrome. Mainly targeting the collagen´s composition in each described part the Picru-sirius Red´s stain under polarized light was used. Among thirty-four penguin hearts, none presented discrepancy in morphology, they were all very similar. The characterization of collagen by Picrusirius Red stain highlighted type 1 collagen in comparison to type 3 collagen in most structures, giving a more reddish appearance in almost of them. In conclusion, the cardiac anatomy of the Magellanic Penguin is similar to that of other birds, with a predominance of type I collagen.(AU)

A Paleocene penguin from New Zealand substantiates multiple origins of gigantism in fossil Sphenisciformes.

One of the notable features of penguin evolution is the occurrence of very large species in the early Cenozoic, whose body size greatly exceeded that of the largest extant penguins. Here we describe a new giant species from the late Paleocene of New Zealand that documents the very early evolution of large body size in penguins. Kumimanu biceae, n. gen. et sp. is larger than all other fossil penguins that have substantial skeletal portions preserved. Several plesiomorphic features place the new species outside a clade including all post-Paleocene giant penguins. It is phylogenetically separated from giant Eocene and Oligocene penguin species by various smaller taxa, which indicates multiple origins of giant size in penguin evolution. That a penguin rivaling the largest previously known species existed in the Paleocene suggests that gigantism in penguins arose shortly after these birds became flightless divers. Our study therefore strengthens previous suggestions that the absence of very large penguins today is likely due to the Oligo-Miocene radiation of marine mammals.

Long necks enhance and constrain foraging capacity in aquatic vertebrates.

Highly specialized diving birds display substantial dichotomy in neck length with, for example, cormorants and anhingas having extreme necks, while penguins and auks have minimized necks. We attached acceleration loggers to Imperial cormorants Phalacrocorax atriceps and Magellanic penguins Spheniscus magellanicus, both foraging in waters over the Patagonian Shelf, to examine the difference in movement between their respective heads and bodies in an attempt to explain this dichotomy. The penguins had head and body attitudes and movements that broadly concurred throughout all phases of their dives. By contrast, although the cormorants followed this pattern during the descent and ascent phases of dives, during the bottom (foraging) phase of the dive, the head angle differed widely from that of the body and its dynamism (measured using vectorial dynamic acceleration) was over four times greater. A simple model indicated that having the head on an extended neck would allow these cormorants to half the energy expenditure that they would expend if their body moved in the way their heads did. This apparently energy-saving solution is likely to lead to greater heat loss though and would seem tenable in slow-swimming species because the loss of streamlining that it engenders would make it detrimental for fast-swimming taxa such as penguins.

Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins.

The total-evidence approach to divergence time dating uses molecular and morphological data from extant and fossil species to infer phylogenetic relationships, species divergence times, and macroevolutionary parameters in a single coherent framework. Current model-based implementations of this approach lack an appropriate model for the tree describing the diversification and fossilization process and can produce estimates that lead to erroneous conclusions. We address this shortcoming by providing a total-evidence method implemented in a Bayesian framework. This approach uses a mechanistic tree prior to describe the underlying diversification process that generated the tree of extant and fossil taxa. Previous attempts to apply the total-evidence approach have used tree priors that do not account for the possibility that fossil samples may be direct ancestors of other samples, that is, ancestors of fossil or extant species or of clades. The fossilized birth­death (FBD) process explicitly models the diversification, fossilization, and sampling processes and naturally allows for sampled ancestors. This model was recently applied to estimate divergence times based on molecular data and fossil occurrence dates. We incorporate the FBD model and a model of morphological trait evolution into a Bayesian total-evidence approach to dating species phylogenies. We apply this method to extant and fossil penguins and show that the modern penguins radiated much more recently than has been previously estimated, with the basal divergence in the crown clade occurring at ∼12.7 ∼12.7 Ma and most splits leading to extant species occurring in the last 2 myr. Our results demonstrate that including stem-fossil diversity can greatly improve the estimates of the divergence times of crown taxa. The method is available in BEAST2 (version 2.4) software www.beast2.org with packages SA (version at least 1.1.4) and morph-models (version at least 1.0.4) installed.

A new fossil from the mid-Paleocene of New Zealand reveals an unexpected diversity of world's oldest penguins.

We describe leg bones of a giant penguin from the mid-Paleocene Waipara Greensand of New Zealand. The specimens were found at the type locality of Waimanu manneringi and together with this species they constitute the oldest penguin fossils known to date. Tarsometatarsus dimensions indicate a species that reached the size of Anthropornis nordenskjoeldi, one of the largest known penguin species. Stem group penguins therefore attained a giant size very early in their evolution, with this gigantism existing for more than 30 million years. The new fossils are from a species that is phylogenetically more derived than Waimanu, and the unexpected coexistence of Waimanu with more derived stem group Sphenisciformes documents a previously unknown diversity amongst the world's oldest penguins. The characteristic tarsometatarsus shape of penguins evolved early on, and the significant morphological disparity between Waimanu and the new fossil conflicts with recent Paleocene divergence estimates for penguins, suggesting an older, Late Cretaceous, origin.

Novel insights into early neuroanatomical evolution in penguins from the oldest described penguin brain endocast.

Digital methodologies for rendering the gross morphology of the brain from X-ray computed tomography data have expanded our current understanding of the origin and evolution of avian neuroanatomy and provided new perspectives on the cognition and behavior of birds in deep time. However, fossil skulls germane to extracting digital endocasts from early stem members of extant avian lineages remain exceptionally rare. Data from early-diverging species of major avian subclades provide key information on ancestral morphologies in Aves and shifts in gross neuroanatomical structure that have occurred within those groups. Here we describe data on the gross morphology of the brain from a mid-to-late Paleocene penguin fossil from New Zealand. This most basal and geochronologically earliest-described endocast from the penguin clade indicates that described neuroanatomical features of early stem penguins, such as lower telencephalic lateral expansion, a relatively wider cerebellum, and lack of cerebellar folding, were present far earlier in penguin history than previously inferred. Limited dorsal expansion of the wulst in the new fossil is a feature seen in outgroup waterbird taxa such as Gaviidae (Loons) and diving Procellariiformes (Shearwaters, Diving Petrels, and allies), indicating that loss of flight may not drastically affect neuroanatomy in diving taxa. Wulst enlargement in the penguin lineage is first seen in the late Eocene, at least 25 million years after loss of flight and cooption of the flight stroke for aquatic diving. Similar to the origin of avian flight, major shifts in gross brain morphology follow, but do not appear to evolve quickly after, acquisition of a novel locomotor mode. Enlargement of the wulst shows a complex pattern across waterbirds, and may be linked to sensory modifications related to prey choice and foraging strategy.

Hidden keys to survival: the type, density, pattern and functional role of emperor penguin body feathers.

Antarctic penguins survive some of the harshest conditions on the planet. Emperor penguins breed on the sea ice where temperatures drop below -40°C and forage in -1.8°C waters. Their ability to maintain 38°C body temperature in these conditions is due in large part to their feathered coat. Penguins have been reported to have the highest contour feather density of any bird, and both filoplumes and plumules (downy feathers) are reported absent in penguins. In studies modelling the heat transfer properties and the potential biomimetic applications of penguin plumage design, the insulative properties of penguin plumage have been attributed to the single afterfeather attached to contour feathers. This attribution of the afterfeather as the sole insulation component has been repeated in subsequent studies. Our results demonstrate the presence of both plumules and filoplumes in the penguin body plumage. The downy plumules are four times denser than afterfeathers and play a key, previously overlooked role in penguin survival. Our study also does not support the report that emperor penguins have the highest contour feather density.

Bone histology in extant and fossil penguins (Aves: Sphenisciformes).

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.

Penguin lungs and air sacs: implications for baroprotection, oxygen stores and buoyancy.

The anatomy and volume of the penguin respiratory system contribute significantly to pulmonary baroprotection, the body O2 store, buoyancy and hence the overall diving physiology of penguins. Therefore, three-dimensional reconstructions from computerized tomographic (CT) scans of live penguins were utilized to measure lung volumes, air sac volumes, tracheobronchial volumes and total body volumes at different inflation pressures in three species with different dive capacities [Adélie (Pygoscelis adeliae), king (Aptenodytes patagonicus) and emperor (A. forsteri) penguins]. Lung volumes scaled to body mass according to published avian allometrics. Air sac volumes at 30 cm H2O (2.94 kPa) inflation pressure, the assumed maximum volume possible prior to deep dives, were two to three times allometric air sac predictions and also two to three times previously determined end-of-dive total air volumes. Although it is unknown whether penguins inhale to such high volumes prior to dives, these values were supported by (a) body density/buoyancy calculations, (b) prior air volume measurements in free-diving ducks and (c) previous suggestions that penguins may exhale air prior to the final portions of deep dives. Based upon air capillary volumes, parabronchial volumes and tracheobronchial volumes estimated from the measured lung/airway volumes and the only available morphometry study of a penguin lung, the presumed maximum air sac volumes resulted in air sac volume to air capillary/parabronchial/tracheobronchial volume ratios that were not large enough to prevent barotrauma to the non-collapsing, rigid air capillaries during the deepest dives of all three species, and during many routine dives of king and emperor penguins. We conclude that volume reduction of airways and lung air spaces, via compression, constriction or blood engorgement, must occur to provide pulmonary baroprotection at depth. It is also possible that relative air capillary and parabronchial volumes are smaller in these deeper-diving species than in the spheniscid penguin of the morphometry study. If penguins do inhale to this maximum air sac volume prior to their deepest dives, the magnitude and distribution of the body O2 store would change considerably. In emperor penguins, total body O2 would increase by 75%, and the respiratory fraction would increase from 33% to 61%. We emphasize that the maximum pre-dive respiratory air volume is still unknown in penguins. However, even lesser increases in air sac volume prior to a dive would still significantly increase the O2 store. More refined evaluations of the respiratory O2 store and baroprotective mechanisms in penguins await further investigation of species-specific lung morphometry, start-of-dive air volumes and body buoyancy, and the possibility of air exhalation during dives.

Central corneal thickness and intraocular pressure in captive black-footed penguins (Spheniscus dermersus).

OBJECTIVE: To determine the central corneal thickness (CCT) by ultrasonic pachymetry and the effect of these values on the measurements of intraocular pressures (IOP) with rebound tonometry (TonoVet(®) ) in a captive flock of black-footed penguins (Spheniscus dermersus). Variations in CCT by age and weight, and variations in IOP by age were compared. ANIMAL STUDIED: Both eyes of 18 clinically normal black-footed penguins (Spheniscus dermersus) were used. PROCEDURE: The IOP was measured by the TonoVet(®) in both eyes of all the penguins. CTT measurements were performed 5 min later in all eyes using an ultrasound pachymeter. RESULTS: The mean IOP values ± SD were 31.77 ± 3.3 mm Hg (range of mean value: 24-38). The mean CCT values were 384.08 ± 30.9 µm (range of mean value: 319-454). There was no correlation between IOP and CCT values (P = 0.125). There was no difference in CCT measurements by age (P = 0.122) or weight (P = 0.779). A correlation was observed (P = 0.032) between IOP values and age. The coefficient of correlation was negative (ρ = -0.207). CONCLUSIONS: Ultrasound pachymetry has shown to be a reliable and easy technique to measure CCT in penguins. No correlation was observed between IOP and CCT values in this study. IOP showed a significant but weak decrease as age increased in the black-footed penguin.