Penguins

Thermoregulation could represent a signicant fraction of the total energy budget of endotherms under unfavourable environmental conditions. This cost affects several traits of the ecology of an organism such as its behaviour, distribution, or life history. Heat produced by muscle contraction during activity can be used to pay for heat loss or thermoregulation in many species (known as “thermal substitution”). This study seeks to unite the effects of temperature, depth, and activity on the energetic costs of endotherm divers using the Magellanic penguin as model species and to evaluate whether penguins may benet from thermal substitution. This species operates under highly variable temperature and depth conditions along its breeding range and provides an ideal natural experiment. A developed thermodynamic model describing foraging activity predicted that the major element affecting heat loss was depth, exacerbated by temperature. Birds liv- ing in colder waters are predicted to be able to minimize costs by executing shallower dives and bene t from ther- mal substitution by swimming faster, particularly during deeper dives. The model was evaluated in two contrasting scenarios: (1) when birds swim near the surface commuting to the foraging areas and (2) when birds dive to depth to forage. Activity data from tags on free-living penguins indicated two of these predictions were apparent; penguins generally travelled faster while commuting at the surface in colder waters, while birds from warmer water colonies dived deeper while foraging. Contrary to predictions, how- ever, penguins swam slower at deeper depths during both descent and ascent phases of foraging dives. These results suggest that penguins may bene t from thermal substitu- tion by swimming faster when birds perform shallow dives commuting to and back from foraging areas, but they pro- vide no evidence of behavioural response (via swimming faster) for thermoregulation when diving to depth to forage. Reasons for this are discussed and include the relevance of prey abundance in 3-d space and maximizing dive duration by conserving oxygen reserves. The way the bird operates will have profound consequences for the energy needed and therefore necessary energy acquisition rates. Expansion of our ndings to other diving endotherms might help explain both global activity patterns and energy ow in ecosystems.

Chinstrap and Adeíie penguins generate considerable pressures to propel their faeces away from the edge of the nest. The pressures involved can be approximated if the following parameters are known: (1) distance the faecal material travels before it hits the ground, (2) density and viscosity of the material, and (3) shape, aperture, and height above the ground of the orificium venti. With all of these parameters measured, we calculated that fully grown penguins generate pressures of around 10 kPa (77 mm Hg) to expel watery material and 60 kPa (450 mm Hg) to expel material of higher viscosity similar to that of olive oil. The forces involved, lying well above those known for humans, are high, but do not lead to an energetically wasteful turbulent flow. Whether a bird chooses the direction into which it decides to expel its faeces, and what role the wind plays in this, remain unknown.

Valuing objects is a distinctive human trait. It is necessary for rational behaviour. Factors that are likely to influence valuations, the difficulties of getting agreements about valuations and the limited perspective of economics as a basis for valuation are discussed generally in this article. Attributes of Otago Peninsula that seem to be valuable and worth conserving are listed and discussed, taking into account possible conflicts in getting maximum value from these attributes. Particular attention is given to the economic value of conserving wildlife species on the Otago Peninsula. As a result of the presence of these species and their use for tourism, expenditure of over $100 million NZ is generated annually in the Dunedin regional economy (directly or indirectly) and that 800-1000 full-time equivalent jobs are created. The economic opportunity cost of this wildlife conservation on the Peninsula is low and that the economic benefits from this conservation are well in excess of the costs involved. When non-use economic values and the social values associated with Otago Peninsula are taken into account, this further adds to the value of conserving this wildlife. While there has been remarkable expansion in wildlife tourism on the Otago Peninsula and its economic impact in the Dunedin region in the last two decades, (especially in the viewing of Yellow-eyed Penguins), difficulties and constraints are emerging that are likely to hamper its future expansion. However, wildlife tourism on the Otago Peninsula will still have a huge economic impact on Dunedin’s regional economy in the future. Consequently, even if assessed solely in terms of their economic value, the wildlife attractions of the Otago Peninsula are well worth conserving. Furthermore, the value of conserving biodiversity on the Peninsula exceeds its touristic and its economic value. Some conservation organizations (such as the Yellow-eyed Penguin Trust), even though not directly involved in tourism, add to its economic value as well as to its social value and promote the highly desired goal of conserving biodiversity.

It is suggested that valuing objects is a distinctive human trait. It is a prerequisite for rational behaviour. Factors that are likely to influence valuations, the difficulties of getting agreements about valuations and the limited perspective of economics as a basis for valuation are discussed generally. Attributes of Otago Peninsula that seem to be valuable and worth conserving are listed and discuss taking into account possible conflicts in getting maximum value from these attributes. Particular attention is given to the economic value of conserving wildlife species on the Otago Peninsula. It is estimated that as a result of the presence of these species and their use for tourism, expenditure of over $100 million annually is generated in the Dunedin regional economy (directly or indirectly) and that 800-1000 full-time equivalent jobs are created. The economic opportunity cost of this wildlife conservation on the Peninsula is low and it is clear that the economic benefits from this conservation are well in excess of the costs involved. When non-use economic values and the social values associated with Otago Peninsula are taken into account, this further adds to the value of conserving this wildlife. While there has been remarkable expansion in wildlife tourism on the Otago Peninsula and its economic impact in the Dunedin region in the last two decades, (especially in the viewing of yellow-eyed penguins) difficulties and constraints are emerging that are likely to hamper its future expansion. However, wildlife tourism on the Otago Peninsula will still have a huge economic impact on Dunedin regional economy. Consequently, even if assessed solely in terms of their economic value, the wildlife attractions of the Otago Peninsula are well worth conserving. It can be argued however, that the value of conserving biodiversity on the Peninsula exceeds its touristic and its economic value. Some conservation organizations (such as the Yellow-eyed Penguin Trust) even though not directly involved in tourism add to its economic value as well as its social value and promote the highly desired goal of conserving biodiversity.

This article updates a paper which I wrote in 1988 about the economic value of biodiversity conservation on the Otago Peninsula and the scope for expanding wildlife tourism there. After outlining different ways to measure the economic importance of wildlife, I use economic impact analysis to measure the current importance of wildlife tourism on the Otago Peninsula. However, it is also pointed out that the activities of bodies such as the Yellow-eyed Penguin Trust and the Department of Conservation have positive regional economic impacts. The specific methods and assumptions used for this economic analysis are outlined. The gross annual turnover of enterprises directly involved in the viewing of wildlife on the Peninsula is of the order of $6.5 million per year and they employ the equivalent of 70 full-time persons. With multiplier or flow-on effects these economic magnitudes are higher. However, the economic impact of wildlife tourism based on the Peninsula is much greater. The presence of wildlife on the Peninsula attracts travellers to the Dunedin region who would otherwise not visit and entices some who would have visited anyway to stay longer. This increases local expenditure on accommodation, food and so on. As a result, it is estimated that an extra $100 million, or so in expenditure occurs in Dunedin’s regional economy and employment is increased by the equivalent of 800-1000 full-time positions. The economic impact of wildlife on Dunedin’s regional economy has increased by more than eleven-fold in the last 20 years.

While growth in tourism on the Peninsula is still possible, it is likely to be at a slower rate than in the past. This is because capacity constraints are being encountered. Furthermore, the future security of the Peninsula’s flagship species is not assured. In addition, problems are emerging where there is free access to areas where wildlife may be seen. It is observed that the opportunity cost of conserving most wildlife on the Otago Peninsula is low, but some conflict may be occurring between wildlife conservation and human uses of marine areas. The paper, however, makes it quite clear that the loss of wildlife on the Otago Peninsula would result in a huge economic loss to the Dunedin region.

The modern zoo has brought about two major advances in the behavioral welfare of their exhibited animals: (a) The use of environmental enrichment to promote naturalistic behaviors and (b) the use of training to improve voluntary husbandry care. Whereas training itself has been talked about as an effective enrichment strategy, little has been done to combine training procedures with enrichment. Typically, enrichment is treated as a trial and error process, where potential enrichment items or procedures are cycled through until successful enrichment is found. The use of shaping or other training techniques has seldom been documented to increase engagement with possible enrichment items or procedures. The following study examined the possibility of combining training and enrichment to produce continued interactions with enrichment devices. Two species of penguin, Magellanic and southern rockhopper penguins, were studied. Two measures were taken: Time spent swimming and contact with enrichment devices. The enrichment devices could be manipulated by placing fish within and hanging out of each device. During baseline sessions, no hits to either device were observed. During training sessions, several hits were recorded when fish were in the devices and overall swimming time increased during these conditions. When baseline was reintroduced without fish in the devices, contact with the enrichment devices rapidly declined and swimming time for the rockhopper penguins decreased. When the devices were reintroduced with fish but without training, the greatest number of enrichment device contacts and the highest percentage of time spent swimming were observed for the rockhopper penguins.

We developed a Hidden Markov mark–recapture model (R package marked) to examine sex‐specific demography in Magellanic Penguins (Spheniscus magellanicus). Our model was based on 33 yr of resightings at Punta Tombo, Argentina, where we banded ~44,000 chicks from 1983 to 2010. Because we sexed only 57% of individuals over their lifetime, we treated sex as an uncertain state in our model. Our goals were to provide insight into the population dynamics of this declining colony, to inform conservation of this species, and to highlight the importance of considering sex‐specific vital rates in demographic seabird studies. Like many other seabirds, Magellanic Penguins are long‐lived, serially monogamous, and exhibit obligate biparental care. We found that the non‐breeding‐season survival of females was lower than that of males and that the magnitude of this bias was highest for juveniles. Biases in survival accumulated as cohorts aged, leading to increasingly skewed sex ratios. The survival bias was greatest in years when overall survival was low, that is, females fared disproportionality worse when conditions were unfavorable. Our model‐estimated survival patterns are consistent with independent data on carcasses from the species’ non‐breeding grounds, showing that mortality is higher for juveniles than for adults and higher for females than for males. Juveniles may be less efficient foragers than adults are and, because of their smaller size, females may show less resilience to food scarcity than males. We used perturbation analysis of a population matrix model to determine the impact of sex‐biased survival on adult sex ratio and population growth rate at Punta Tombo. We found that adult sex ratio and population growth rate have the greatest proportional response, that is, elasticity, to female pre‐breeder and adult survival. Sex bias in juvenile survival (i.e., lower survival of females) made the greatest contribution to population declines from 1990 to 2009. Because starvation is a leading cause of morality in juveniles and adults, precautionary fisheries and spatial management in the region could help to slow population decline. Our data add to growing evidence that knowledge of sex‐specific demography and sex ratios are necessary for accurate assessment of seabird population trends.

Penguins are considered among the most popular animals for zoo and aquarium visitors to observe. Swimming is considered a desirable activity, both for the visitor experience and the welfare of the penguins. However, little is known about the amount of time exhibited penguins spend swimming, or how such swimming is related to regular feeding events. We examined the effects of introducing live prey in the form of trout on 22 Humboldt penguins living at the Woodland Park Zoo. Of primary interest was how the live feeds changed (1) daily and hourly swimming activity, and (2) variability in enclosure use. We hypothesized that the live feedings would increase swimming activity prior to and during the delivery of the live trout, as well as create an overall increase in total swimming activity for live feed days compared to non-live feed days. We also predicted that the penguins would be more likely to use the entire exhibit around these live feeds, since they are likely to chase fish throughout the exhibit. Penguins did show an increase in swimming activity in the hour prior to and during the live feed, with a small decrease in swimming activity following the live feed when compared to non-live feed days. There was also a more than 30% increase in the total swimming activity for live feed days when compared to all other non-live feed days. In addition, a single measure of variability in enclosure use (entropy) showed greater overall enclosure use for the live feed days compared to the non-live feed days. These results demonstrate that live fish can be a useful way of enriching the behavioural welfare of Humboldt penguins.

1. Nonlinear, parametric curve-fitting provides a framework for understanding diverse ecological and evolutionary trends (e.g. growth patterns and seasonal cycles). Currently, parametric curve-fitting requires a priori assumptions of curve trajectories, restricting their use for exploratory analyses. Furthermore, use of analytical techniques [nonlinear least-squares (NLS) and nonlinear mixed-effects models] for complex parametric curves requires efficient choice of starting parameters.


2. We illustrate the new R package FlexParamCurve that automates curve selection and provides tools to analyse nonmonotonic curve data in NLS and nonlinear mixed-effects models. Examples include empirical and simulated data sets for the growth of seabird chicks.


3. By automating curve selection and parameterization during curve-fitting, FlexParamCurve extends current possibilities for parametric analysis in ecological and evolutionary studies.

Many birds can visualise parts of the ultraviolet
(UV) range; consequently, the diversity of colours
observed by birds is generally greater than humans
can perceive. The colouration of feathers is diverse
both between and within bird species. Feather colour
can result from either pigments or structure—
i.e. when light refracts off of the physical structure
of the feather—or a combination of both. In the little
penguin (Eudyptula minor), also known as the little
blue penguin, non-iridescent (β-keratin) nanostructures
result in their characteristic blue colour. As little penguins are categorised into two clades (i.e.
Australia/southeast New Zealand clade and the New
Zealand-only clade), we compared different colour
characteristics of feathers by sex and clade. Using a
recently designed spectrometry instrument to reliably
quantify structural colours, we analysed two to
three feathers from each individual. We also used
feather-based DNA to determine sex and clade for
each individual. Preliminarily results suggest that
less than 30% of feathers analysed have a characteristic
UV/Blue hue (300–510 nm) whereas over
70% were considered simply shiny (hue approx.
700 nm). Of those individuals demonstrating peaks
in the UV/Blue range in any of their feathers
sampled (124 of 259 birds), we found a significant
difference between clades for UV chroma (proportion of brightness in UV range), blue chroma
(proportion of brightness in blue range), and hue
(wavelength of maximum reflectance peak). We
detected no difference between sex for UV chroma,
blue chroma, or hue, but we did detect a significant
difference between male and female feather maximum
brightness (area under the maximum spectral
curve). These differences may be related to sexual
dimorphism and clade differentiation within the
species, but the biological significance of these
differences is unclear. We are expanding this study
to include feather samples from more birds of both
clades as well as analysing the colour pattern on the
dorsal side of the flipper, which may also vary by sex
and clade.

Improving husbandry parameters of Bird of Paradise, utilizing state of the art medical equipment to evaluate the health of the Little Penguins and employing aviculture techniques developed at the Bronx Zoo to augment field research are all part of our ongoing work to enhance the conservation of birds. We are constantly striving to improve animal husbandry, the processes involved in the feeding, propagation, housing and health care of the birds at the WCS to ensure the maximum benefits to those animals. The impetus for these efforts is the establishment of viable populations of birds in conservation care that contribute to the improved knowledge and conservation of wild bird populations. Programs, within the WCS's Bronx Zoo, directly influence conservation strategies of threatened bird species as we collaborate with IUCN SSC, governmental agencies and other NGO's to achieve our goals. Management of in-situ populations of birds has proven to increase wild populations and many of those protocols were developed in conjunction with the populations of birds in managed programs. Please read further to discover a wide range of activities that are underway to improve avian husbandry and in turn affect the health of our planet by improving the resilience of wildlife and wild places.

The penguins (Aves: Sphenisciformes) are the most derived group of diving seabirds. Exclusive to the Southern Hemisphere, ten of the seventeen extant species can be found in South America. South America has been recognised as one of the richest areas with fossil penguins worldwide, but until the late twentieth century this record was limited to the Argentinian Patagonia. Only at the beginning of this century were the first species from southern Peru and northern Chile described. Currently, seventeen palaeospecies have been described for South America, with a chronostratigraphic range from Middle Eocene to Late Pliocene. Traditionally the study of fossil penguins has been made on the basis of isolated specimens, with the humerus and tarsometatarsus being the most widely used elements for the typification of species. However, recently the number of skulls and partial skeletons discovered in South America has significantly increased, facilitating their interpretation and use in phylogenetic analyses. This work summarizes the fossil record of Sphenisciformes on the continent, based on type specimen revisions and the compilation of bibliographic data.

The present review aims to improve the scope and coverage of the phylogenetic matrices currently in use, as well as explore some aspects of the relationships among Paleogene penguins, using two key skeletal elements, the humerus and tarsometatarsus. These bones are extremely important for phylogenetic analyses based on fossils because they are commonly found solid specimens, often selected as holo− and paratypes of fossil taxa. The resulting dataset includes 25 new characters, making a total of 75 characters, along with eight previously uncoded taxa for a total of 48. The incorporation and analysis of this corrected subset of morphological characters raise some interesting questions consider− ing the relationships among Paleogene penguins, particularly regarding the possible exis− tence of two separate clades including Palaeeudyptes and Paraptenodytes, the monophyly of Platydyptes and Paraptenodytes, and the position of Anthropornis. Additionally, Noto− dyptes wimani is here recovered in the same collapsed node as Archaeospheniscus and not within Delphinornis, as in former analyses.

A comparison between the Paleogene and Neogene marine assemblages in South America suggests a major turnover among penguins by the late Miocene, with the disappearance of the stem Sphenisciformes and the rise of the modern penguins. However, isolated specimens attributed to Palaeospheniscus from the late Miocene of Chile suggest that stem and crown taxa coexisted until the end of the Miocene. To clarify the stratigraphic range of Palaeospheniscus and determine how long the stem Sphenisciformes persisted in South America, the present study offers a revision of its putative records in Chile (Bahía Inglesa and Coquimbo formations), Peru (Chilcatay Formation), and South Africa (Ysterplaats site). In order to reliably discriminate isolated humeri of Palaeospheniscus from Spheniscus, twelve potential discriminating characters were identified and evaluated. Despite the existence of a wide range of variability and overlap, these humeri can be discriminated using large sets of characters, such as a transverse ligament sulcus connected with the capital incisure by a sulcus, or the lack of the proximal ligament pit in Palaeospheniscus. The putative record of Palaeospheniscus in the late Miocene–Pliocene of Chile and early Pliocene of South Africa is discarded, restricting its stratigraphic range to the early and middle Miocene of Argentina and the middle Miocene of Peru. This reduces the biochron of the genus from 18 Ma to a maximum of 11 Ma and suggests that the middle Miocene was a crucial time for the establishment of modern seabirds communities in general and penguins in particular.

Land-breeding marine animals such as penguins, flying seabirds and pinnipeds are important components of marine ecosystems, and their abundance has been used extensively as an indication of ecosystem status and change. Until recently, many efforts to measure and monitor abundance of these species’ groups have focussed on smaller populations and spatial scales, and efforts to account for perception bias and availability bias have been variable and often ad hoc. We describe a suite of new methods, technologies and estimation procedures for costeffective, large-scale abundance estimation within a general estimation framework and illustrate their application on large Ade´lie penguin populations in two regions of East Antarctica. The methods include photographic sample counts, automated cameras for collecting availability data, and bootstrap estimation to adjust counts for the sampling fraction, perception bias, and availability bias, and are
applicable for a range of land-breeding marine species. The
methods will improve our ability to obtain population data
over large spatial and population scales within tight logistic, environmental and time constraints. This first application of the methods has given new insights into the biases and uncertainties in abundance estimation for penguins and other land-breeding marine species. We provide guidelines for applying the methods in future surveys.

During the last decade, new Neogene fossil assemblages from South America have revealed important clues about the evolution of seabird faunas in one of the major upwelling systems of the world: the Humboldt Current. However, most of this record comes from arid Northern Chile and Southern Peru and, in consequence, our knowledge of the evolutionary history of seabirds in the temperate transitional zone is negligible. A new Late Pliocene assemblage of fossil birds from the coastal locality of Horcon in Central Chile offers a unique opportunity to fill this gap. Isolated bones of a medium-sized penguin are the most abundant bird remains. Morphological and cladistic analyses reveal that these specimens represent a new species of crested penguin, Eudyptes calauina sp. nov. Eudyptes is a penguin genus that inhabit temperate and subantarctic regions and currently absent in central Chile. Additionally, a partial skeleton of a small species of cormorant and a partial tarsometatarsus of a sooty shearwater have been identified. The Horcon fossils suggest the existence of a mixed avifauna in central Chile during the Pliocene in concordance with the latitudinal thermal gradient. This resembles the current assemblages from the transitional zone, with the presence of species shared with Northern Chile and Southern Peru and a previously unrecorded penguin currently absent from the Humboldt System but present in the Magellanic region. Comparison of Pliocene seabird diversity across the Pacific coast of South America shows that the Horcon avifauna represents a distinctive assemblage linking the living faunas with the Late Miocene ones. A comparison with the fossil record near the Benguela Current (west coast of southern Africa) suggests that the thermic gradient could play an important role in the preservation of a higher diversity of cold/temperate seabirds in the Humboldt Current.

Sustainability has emerged as an important field of research with an emphasis on
studying tourism impacts and defining the limits of growth in tourism. However, the
conceptual foundation of sustainability and the limits to growth have been perceived
as vague and their implementation in tourism operations has emerged as a difficult
task. This research note aims to present two different perspectives on sustainability
by utilizing an experimental pilot example (yellow-eyed penguins) to demonstrate
how different theoretical conceptualizations are manifested in nature-based tourism
operations.

A group of zoo-housed little penguins (Eudyptula minor) was diagnosed with mycobacteriosis. While undergoing multidetector computed tomography (MDCT) imaging for an unrelated research project, pulmonary lesions were detected in multiple individuals. In general, birds appeared healthy and free of outward signs of disease. After the loss of three individuals, polyclonal mycobacterial disease due to Mycobacterium avium-intracellulare complex was confirmed. Surviving birds were treated with rifampin (45 mg/kg), ethambutol (30 mg/kg), clarithromycin (10 mg/kg), and enrofloxacin (30 mg/kg) compounded into a single capsule administered once a day in food. After 3 mo of therapy, MDCT imaging documented a decrease in nodule size and number in all remaining birds, with further improvement documented after 13 mo of treatment. MDCT imaging was invaluable for diagnosing disease, documenting disease progression over time, and assessing response to therapy. Early initiation of therapy before the development of outward signs of disease led to resolution of mycobacterial pulmonary lesions in multiple penguins. Mycobacterial disease in this group of little penguins, as well as previously published reports, suggests that the species is at increased risk for developing mycobacteriosis

No-take zones may protect populations of targeted marine species and restore the integrity of marine ecosystems, but it is unclear whether they benefit top predators that rely on mobile pelagic fish. In South Africa, foraging effort of breeding African penguins decreased by 30 % within 3 months of closing a 20-km zone to the competing purse-seine fisheries around their largest colony. After the fishing ban, most of the penguins from this island had shifted their feeding effort inside the closed area. Birds breeding at another colony situated 50 km away, whose fishing grounds remained open to fishing, increased their foraging effort during the same period.". This demonstrates the immediate benefit of a relatively small no-take zone for a marine top predator relying on pelagic prey. Selecting such small protected areas may be an important first conservation step, minimizing stakeholder conflicts and easing compliance, while ensuring benefit for the ecosystems within these habitats

Se presentan los primeros restos diagnósticos asignables a Spheniscus megaramphus procedentes del Miembro Bonebed de la Formación Bahía Inglesa (Mioceno tardío), consistentes en tres rostros aislados pertenecientes a la colección del Museo Paleontológico de Caldera, Chile. Se rectifica la antigüedad de la especie en el Perú, previamente asociada al Mesiniano (6 Ma) y actualmente referida al Tortoniano (9.5-9 Ma). El presente estudio confirma la existencia de esta especie en Chile, ampliando el rango de incidencia de la especie hasta los 27º S durante el Mioceno tardío. ABSTRACT The first diagnostic remains assignable to Spheniscus megaramphus coming from the Bonebed Member of the Bahia Inglesa Formation (Late Mioceno) are presented. These consist of three isolated rostrums from the collection of the Museo Paleontologico de Caldera, Chile. The antiquity of the species is rectified in Peru previously associated to the Mesinian (6 My) and at the moment referred to the Tortonian (9.5-9 My). The present study confirms the existence of this species in Chile enlarging the range of incidence of the species until the 27º S during the Late Miocene.

Sex-biased mortality can increase the risk of extinction of threatened populations.  Numbers of the Endangered African penguins Spheniscus demersus are decreasing rapidly and the smaller size of females associated with their higher foraging effort during the breeding season may put them more at risk than males. Using records from 2004-2012 from a rehabilitation centre in South Africa, we investigated the sex ratio of African penguins that died at the centre within age-classes (adult, juvenile and chick) and causes of death (starvation, injury or oiling). The large majority of adult and juvenile birds at the centre died following starvation, rather than from injuries or oiling, and most of them (>60%) were females, while mortality of sexes was similar among chicks. Oiled birds received at the centre had high survival and release rates, hence sex-biased mortality could not be tested for oiled birds. Female-biased mortality in African penguins could skew adult sex ratio, which may further accelerate the rapid rate of decline of this endangered species.