Blog articles on fun facts and interesting information written by our penguin experts.

Malaria and the African Penguin

african penguin

Malaria and the African Penguin

By Megan Spofford

Africa’s only species of penguin is in serious danger of extinction. Threats to the species have come to a head from guano and egg harvesting (that took place until the 1930s), competition for food resources due to increasing ocean temperatures and humans overfishing, invasive species, and oil spills. African Penguins are listed as endangered on the IUCN Red List. An assessment of the species was last done in August of 2018 that concluded the population trend of roughly 50,000 adult individuals was still in a major decline. The assessment goes on to state that the “…trend currently shows no sign of reversing, and immediate conservation action is required to prevent further declines.” ¹

All penguins are susceptible to malaria

Because of this warning, many institutions have pledged to take part in rehabilitation and breeding efforts to save the African Penguin from extinction. Unfortunately, all penguins are incredibly susceptible to malaria and the disease has a higher occurrence rate in penguins under human care than those in the wild. The disease was first noted in African Penguins in 1968. Specific action against avian malaria must be taken to successfully contribute to conservation efforts that occur in human care for this species.

Details about malaria in penguins and other birds

P. relictum in african penguin
Blood smear showing P. relictum infection (stained purple) within red blood cells. Public Domain.

Malaria itself is interesting, because it is a parasitic disease caused by Plasmodium protozoans of which there are 7 different species that occur in penguins. The parasite is injected into the host organism (penguin) during a blood meal from a vector (mosquito) after it has been picked up from a reservoir (wildlife in close vicinity).

Recent studies involving blood smear analyses from wild penguins that go into rehabilitation facilities showed that roughly ⅓ of the birds admitted were positive for malaria-fighting antibodies, meaning they had already had a malaria infection at some point in their life. These initial samples were taken upon admittance to the facility, so data could determine whether subsequent samples showed presence of new or original disease infection.

Weekly blood samples were then taken of all African Penguins and data shows that malaria infection acquired while being rehabilitated most likely resulted from infection by a different species of Plasmodium than the one that had showed up in some of the initial testing. 

There is a concern at this point that when a penguin infected with the new species of Plasmodium is integrated back into its original colony, it then becomes a reservoir for that new species to infect the other members.

Because symptoms of infection do not typically appear until the disease has progressed to the point that it is difficult to successfully treat, diagnostic testing of blood samples while still in human care can be routinely done to check for infection, and should be utilized for early detection. 

Treatment for malaria in penguins

Early treatment with anti-malarial prophylactics such as chloroquine and primaquine have proven to be successful in limiting symptoms, but not curing the disease. With African Penguin numbers declining, we could hardly imagine the idea of using them as a human-disease model, but malaria affects penguins and humans in the same manner, so these drugs that have worked for penguins have also worked for humans. 

During summer and early fall months, when temperatures are most conducive to mosquito activity, prophylactics can also be given as a preventative measure. Chronic use of prophylactics is not suggested because it is not cost effective during cooler months, but most importantly because it can limit the development of a penguin’s natural immunity.

Evidence shows that once an African Penguin has been infected and overcome a malaria infection, the antibodies are able to better fight off reinfection. (Females may even pass this immunity to offspring!) Vaccines, which introduce infections to the body in small enough amounts that do not cause sickness, but do signal antibodies to be created, have shown effectiveness against mortality from malaria if administered yearly. 

african penguin

Prevention of avian malaria in penguins is the most important key

Above all, the best form of protection is prevention. Simply enclosing housing facilities with mosquito-proof netting and using mosquito repellants around the facility can drastically reduce the chance that a mosquito will infect an African Penguin with malaria at all.

As research uncovers more information about how to better prevent and/or treat malaria infections in African Penguins being cared for by humans, this conservation problem will cease to exist. And wouldn’t that just be ideal!?

* Note: African Penguins are not the only species of penguin who are capable of contracting malaria… they all are! Especially those that spend time in sub-tropical locations, or anywhere mosquitoes exist.

Did you know penguins could get malaria? Let us know what you learned from this blog. Also, please help us continue to learn more about penguins by donating to Penguins International.

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  1. BirdLife International 2018. Spheniscus demersus. The IUCN Red List of Threatened Species 2018: e.T22697810A132604504. Downloaded on 22 August 2019.
  2. Botes, Annelise, Thiart, Hanlie, Parsons, Nola J., & Bellstedt, Dirk U.. (2017). Conservation implications of avian malaria exposure for African penguins during rehabilitation. South African Journal of Science, 113(7-8), 1-8.
  3. Dashiell, N. (2019, June 25). Animal Research and the Fight Against Malaria – FBR. Retrieved from
  4. M. L. Grilo, R. E. T. Vanstreels, R. Wallace, D. García-Párraga, É. M. Braga, J. Chitty, J. L. Catão-Dias & L. M. Madeira de Carvalho (2016) Malaria in penguins – current perceptions, Avian Pathology, 45:4, 393-407, DOI: 10.1080/03079457.2016.1149145

Penguin Postcards and Port Lockroy

Gentoo Penguins at Port Lockroy

Penguin Postcards and Port Lockroy

by Jodie King

Have you ever gotten to see a penguin in the wild? Have you been on a penguin tourism trip? With the ability to book cheap flights at the touch of a key to beautiful destinations across the world, our access to wildlife is vast and there are very few places left on earth which are untouched by humans. However, not all trips are as glossy as a travel magazine or the serene shots of Instagram, and many horror stories exist surrounding ecotourism. Such as tigers sedated to pose for photographs and tourists allowed to ride endangered elephants.

But have you ever thought about the impact of tourism on wild penguins? There is very little in the media which discusses the impact of our holidays on penguins out in the wild (or on naturalistic reserves). Despite this, there are many articles which list the top destinations to see penguins. For the everyday person trying to book a holiday of a lifetime, it can be confusing to differentiate between activities involving animals that are harmless, those which contribute financially to their conservation, and those that are actually having a negative impact. 

Penguin tourism and Port Lockroy

One of the examples that comes to my mind is a post office at Port Lockroy in Antarctica, which has been televised in recent years and is surrounded by a huge colony of Gentoo Penguins. The first base in the harbour was built in 1944 and has been used to accommodate explorers and scientists. However, the Gentoo Penguins did not arrive until 19851. Guests are allowed to visit the museum and base, and also get to see the penguins in their natural habitat. The UK Antarctic Heritage Trust leaves half the island off limits to visitors, to allow breeding success of the penguins to be monitored to investigate the impact of tourism on the colony. Despite having up to 18,000 visitors per season the results reveal no impact on the breeding success of the penguins.

Gentoo Penguins at Port Lockroy
Gentoo Penguin

Penguin tourism does affect some penguins negatively

Yellow-eyed Penguin
Yellow-Eyed Penguin

Some reserves however, are seeing an impact, and not for the better. Katiki Point in New Zealand has a reserve which is home to Yellow-eyed Penguins. They have found saddening results that only half the number of chicks were raised in 2014/15 in areas with visitors, compared to colonies close by which were not impacted by tourism2. Nests which were within 10m of the public paths were the most affected. They found that chicks were requiring help with fledging the nest and they were having to assist parent penguins with their annual moult thought to be caused by stress from visitors.

Other research has found similar negative consequences for Yellow-eyed Penguins which are exposed to tourism. The presence of people reduces the amount of time Yellow-eyed Penguins spend preening3, which is really important to help keep their feathers in good condition to maintain the penguin’s ability to stay dry and protected from the elements. Additionally, despite the use of a distance rule at reserves, the space between a penguin and person is strongly related to disturbance behaviour,3 indicating that perhaps 5m is still too close for comfort.

People also come with baggage. Other things need to be considered with tourism in addition to the people themselves. The potential for the spread of pathogens from person to penguin, and the presence of artificial light, so penguins can actually still be viewed when it’s dark are just two of the factors that need to be considered. But scientists are already on it! Surprisingly, they have found that the introduction of artificial light does not detrimentally impact Little Penguins returning to shore each night and in fact the penguins seemed to prefer paths that were well lit4. Regarding disease, in 2005 research looking for infection found no pathogens from humans in eight different bird species in Antarctica5. So hopefully this will continue to be the case.

Not all penguin tourism has effects on penguins

Thankfully, most penguin tourism destinations enact strict rules where visitors are concerned, requiring guests to stay a certain distance away from the birds at all times, keeping disturbance to a minimum, and evoking firm no contact regulations. New methods are constantly being implemented to try and improve tourism with penguins. Philip Island Reserve in Australia who are fortunate enough to witness the Little Penguins returning to the beaches each night5 are continuously improving their facilities. After seeing the impact people were having on the penguins, the reserve has built underground viewing areas and large seating areas a short distance away from the penguins, to try and minimise the impact on the birds, whilst still allowing visitors to get close to the action. 

But is this the best solution? Making our interactions with wildlife much more regimented could mean fewer negative consequences, whilst making natural reserves far less natural for the birds themselves. Or are there perhaps other options we are yet to explore?

Do you want to travel to see penguins? If so, has this helped you decide where to go? Also, please help us continue to learn more about penguins by donating to Penguins International.

Check out some of our other blogs, too:

Useful references: 



3 FRENCH, R., MULLER, C., CHILVERS, B., & BATTLEY, P. (2019). Behavioural consequences of human disturbance on subantarctic Yellow-eyed Penguins Megadyptes antipodes. Bird Conservation International, 29(2), 277-290. doi:10.1017/S0959270918000096

4 Jonas BonnedahlTina BromanJonas WaldenströmHelena PalmgrenTaina Niskanen, and Björn Olsen “In Search of Human-associated Bacterial Pathogens in Antarctic Wildlife: Report from Six Penguin Colonies Regularly Visited by Tourists,” AMBIO: A Journal of the Human Environment 34(6), 430-432, (1 August 2005).

5 Rodríguez, A., Holmberg, R., Dann, P., & Chiaradia, A. (2018). Penguin colony   attendance under artificial lights for ecotourism. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology, 329(8-9), 457-464.


Penguins and Their Chicks: Super-Parents

King Penguin and its chick

Penguins and Their Chicks: Super-Parents

by Emma Williams

A common misconception about penguin parenting, instigated by polar explorers in the 1960s, was that penguins regularly deserted their chicks. It was believed that they deliberately starved their chicks in order to force them to leave the breeding colony. This was founded on observations that the chicks of King Penguins (Aptenodytes patagonica) weighed more than adult birds and lost this weight prior to fledging. The desertion theory was subsequently generalised to other penguin and seabird species such as Albatrosses1. 

Reproductive effort is a balance between the benefit of increasing offspring survival and costs, including mortality, of the parent. Any increase in parental care must therefore be finely weighed against the price paid. Fundamentally, this boils down to the harsh question: Are they worth feeding?

There are many parenting strategies for successful breeding

Rabbits often produce one litter a month, turtles yield hundreds of hatchlings and provide no parental care, Robins lay 3-4 clutches per year, about half of the chicks do not survive. Penguins tend to go for the opposing parenting strategy of “putting all your eggs in one basket”. And often there is only one egg. 

Far from deserting their young, penguins are super-parents. Compared with most sea-birds, penguins have a very long pre-fledge duration; from 56 days in the Adelie Penguin (Pygoscelis adeliae) to a staggering fifteen months in the King Penguin. Penguin parental care can be divided into two periods. During the guard phase, penguin parents brood the chicks intensively, the female often returns to the sea to forage during this period, which can last up to 37 days. In the next phase the chicks form tight groups, or creches as they’re called1. 

For King Penguin parents, each fledgling represents a huge investment. They first breed when they are around three years old. The parents spend one summer and two winters raising their young. The chicks’ weight loss, as observed by the explorers, can be explained by fluctuations in food availability. King Penguin chicks pile on the calories, lay down fat deposits and balloon to their maximum weight at four months old. A second peak occurs at around ten months. This enables them to survive long periods without food when their parents are foraging. King Penguins are serially monogamous with both parents sharing all hatching and rearing duties. They cannot raise a chick every year which explains why eggs as well as quite large chicks can be seen at the same time in King Penguin colonies2

Penguin parents are even featured in movies

The astonishing parental care provided by the Emperor Penguin (Aptenodytes forsteri) has been popularised by films such as March of the Penguins and Warner Brothers’ Happy Feet. The female lays a single egg that is incubated by the male during the long Antarctic winter. A great deal of the 65-day incubation period is spent in darkness. Standing for weeks, balancing an egg on their feet, with no food, in temperatures down to minus 40 Celsius must be the epitome of good parenting. The female devotes this time to replenishing her food reserves in the open sea. On her return, these penguin parents take turns foraging at sea and caring for the chick in the colony. They are truly long-distance commuters taking foraging trips of one to three weeks averaging over 650 km per trip. They go to great lengths, literally and metaphorically, to care for their young.

The chicks need to gain significant reserves, particularly in the month prior to fledging. A study looking at the diving behaviour of adult Emperor Penguins provisioning chicks during this period, documented some record-breaking feats. They are the deepest diving of all the penguins with dives of over 500m logged. Many dives were well over ten minutes in duration, with the average number reaching more than 200 per day3.

As if this wasn’t enough, Emperor Penguin parents face an additional challenge in successfully rearing a chick: They breed on seasonal sea-ice. The chicks therefore need to fledge before the ice melts in mid-to-late summer. It truly is a race against time.

Penguin chicks hatch into some of the harshest conditions on earth. To survive independently they need to be able to hunt and capture food, often on their first-ever trip to the sea. To do this they need to be strong, fast and capable. The Gentoo Penguin (Pygoscelis papua) manages this challenge in a unique way. A study found two behaviours not shown in other Pygoscelis species: Delayed dispersal of young, and extended parental provisioning4.

Gentoo Penguin feeding its chick

Whilst Adelie Penguins and Chinstrap Penguins (Pygoscelis antarcticus) fledge in a mass exodus on their first trip to the sea, Gentoo chicks take their time and several practice trips. Most Gentoos first go to sea at 70 days old and make approximately five trips in a fortnight before fledging. What is so special about Gentoo Penguins is that they continue to feed their chicks during this period. It is likely that this extended provisioning helps the chicks to hone their hunting skills before fully fledging4.

With extensive pre-fledge periods, long-distance commutes, deep dives, and comprehensive parental provisioning in harsh conditions — often to raise a single chick — penguins justly deserve the accolade “super-parents”.

Do you love the super-parenting of penguins? Let us know! Also, please help us continue to learn more about penguins by donating to Penguins International. We more than appreciate your support!

You can also read more about penguins in the following blogs:

  1. Burger, J. The ‘Desertion Period’ in Seabirds. Proceedings of the Colonial Waterbird Group, vol. 3, 1980, pp. 16–26. JSTOR,
  3. Kooyman, G.L. &. Kooyman, T.G. Diving Behavior of Emperor Penguins Nurturing Chicks at Coulman Island, Antarctica, The Condor: Ornithological Applications, Volume 97, Issue 2, 1 May 1995, Pages 536–549,
  4. Polito, M.J. & Trivelpiece, W.Z. Mar Biol (2008) 154: 231.

King Penguins and Their Vocalizations

King Penguin Facts and Information

King Penguins and Their Vocalizations

by Sian Liversage

Many people have seen the animated movie “Happy Feet,” which is about a young Emperor Penguin chick with a terrible singing voice. Well, penguin voices in the real world are far more vital than people may realise. And although the movie is about Emperor Penguins, I want to talk to you about King Penguins and how their voices are key to the survival of their species.

Facts About King Penguins

First, a few facts about these great birds. King Penguins are the second-largest species of penguin (85-95cm tall), they have a white belly, a silver-grey back, a black head and an obvious striking patch of orange-gold feathers on their neck. They live on vegetated margins in regions of the sub-Antarctic and the Falklands.

During breeding season, they can form vast colonies on snow-free land near the sea, allowing them to forage for food all year around. Colonies vary in size, from 30 birds to tens of thousands, and divided between breeders and non-breeders. Despite the pressure for space and resources, there is very little antagonistic behaviour shown between individuals within the colony. Navigation can be challenging in these large densely populated colonies because of the large number of individuals obstructing the locally available cues.

King Penguins Can Navigate Their Way Through a Colony of Thousands of Birds

When it comes to breeding, both parents help with incubating eggs and brooding chicks, which takes around 9 months for the chick to fledge. Chicks learn the calls of their parents within the first month of their lives, something which proves to be essential. As the chick gets older, the chick is left alone for long periods of time in crèches while its parents forage for food, and on their return, parents can recognise their chick’s calls. Despite the chaos of navigating their way through the colony, these penguins can pinpoint the call of their partner or chick amongst thousands of calling individuals, much in the same way people hear their names in a hub of conversation. This phenomenon is known as the “cocktail party” effect.

King Penguin Facts and Information
King Penguin. Thousands of adults and chicks in a large colony. St Andrew Bay, South Georgia, January 2016. Image © Rebecca Bowater by Rebecca Bowater FPSNZ AFIAP

Despite many studies that are devoted to King Penguin research, there are still many unanswered questions regarding acoustic communication, but in more recent studies, more has been uncovered.

Understanding King Penguin Vocalizations

Researchers conducted playback experiments to truly test their abilities and understand these complex calls1. The experiment involved manipulating different parts of King Penguin’s calls to determine exactly how they recognise each other. Results showed that chicks responded only to the bass frequencies, which travel effectively through a wall of intervening bodies. The first quarter of a second of the parent’s call is enough for the chicks to recognise its parents. The parents will continue to make the call every few seconds. When they are within approximately 11m (36ft) away, the chick will recognise and localise the call. This ability to recognise calls is essential for each breeding pair to successfully raise their chick to adulthood.

Watch the video to hear what King Penguins sound like in the wild. They’re amazingly loud when there are thousands of them together in one colony!

King Penguin Vocalizations May Also Be Different Between Males and Females

Not only has vocalization been discovered as an important signal for individuals, but also for sex recognition. In many penguin colonies it has proven difficult for scientists to determine the sex of the individuals, where females and males are often monomorphic in their external morphology (meaning that they look the same), so there has been limited data on these cues. Another recent study has revealed that King Penguins can be sexed with an accuracy of 100% based on a sex-specific syllable pattern in their vocalizations2. To put this into perspective, a measurement of the bill length of a King Penguin can be sexed with an accuracy of only 79%. These findings may help towards not only understanding how King Penguins choose their mates, but also allows a cost-effective, non-invasive technique for researchers to determine the sex of King Penguins in the field. So future research on King Penguins may have just gotten a whole lot easier.

All in all, given the highly vocal nature of these birds, it is not surprising that chicks learn their parents’ calls within the first month, and adults can be differentiated according to their sex-specific syllable pattern. Based on the current research, it can therefore be concluded that they rely on their vocal signals to recognise each other amongst thousands, therefore making it vital to their survival. Finally, they also present a very interesting system for testing soundscape orientation, an idea that I am sure researchers are already underway in testing.

Did you know about the voices of penguins? Share your thoughts with us, leave us a comment. Also, please help us continue to learn more about penguins by donating to Penguins International.

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  1. Uhlenbroek, C. 2008. Animal life: the definitive visual guide to animals and their behaviours. P431-459
  2. Hannah J. Kriesell, Thierry Aubin, Víctor Planas-Bielsa, Marine Benoiste, Francesco Bonadonna, Hélène Gachot-Neveu, Yvon Le Maho, Quentin Schull, Benoit Vallas, Sandrine Zahn, Céline Le Bohec. Sex identification in King Penguins Aptenodytes patagonicus through morphological and acoustic cues. Ibis, 2018
  3. Anna P. Nesterova, Jules Chiffard, Charline Couchoux, Francesco Bonadonna Journal of Experimental Biology 2013 216: 1491-1500

Bumblefoot in Penguins

bumblefoot in penguins

Bumblefoot in Penguins

by Louise Chiverton

“Bumblefoot” in penguins probably sounds like the cutest disease you could suffer from. Just the word “bumble” makes a person think of some confused little bee which keeps bumping into a window or getting blown off course by the tiniest gust of wind. Unfortunately, the condition is a little more gruesome than the name suggests, so prepare yourselves, because it’s going to get grisly.

Penguins are one of the most popular avian species to be kept in captivity worldwide. Whether you agree or disagree with captive collections, the benefit of these colonies is that we can learn more about the species – their lifecycles, their biology and ultimately how they are impacted by global changes. But by keeping animals in captivity we begin to see an array of new hurdles to overcome; collectively known as “husbandry induced diseases,” and this is where bumblefoot comes into play. Penguins are relatively simple animals to keep due to their small size, basic dietary needs, and they’re easily entertained, spending hours chasing the reflections from a glitter ball. However, they’re not well adapted for the sedentary lifestyle of captivity. 

What is bumblefoot in penguins?

Bumblefoot, or ulcerative pododermatitis refers to the inflammation of the footpad which can be fatal if left untreated. While the primary, blister-like lesions on the base of the foot are not seriously detrimental to overall health, they render the animal susceptible to secondary infection, leading to chronic lameness or septicaemia (Tolpinrud et al., 2017). Warning signs of bumblefoot include limping, lopsided posture to avoid putting pressure on the infected foot and lying down for extended periods of time.  The condition has been identified in a variety of species including rodents, raptors and in rare cases humans (Stransky et al., 2016); each exhibiting the hot, painful lesions on the sole of the foot. However heavy bodied birds which spend more time on foot than in flight appear the most affected taxa.

bumblefoot in penguins
Bumblefoot lesions on the foot of a captive penguin. Image by Louise Chiverton

So, what’s causing bumblefoot? If you think about blisters on a human foot, it’s usually due to friction between the shoe and heel and you’ve probably been walking for a long time (or maybe you’ve just got really sweaty feet!). These factors are similar to those affecting captive penguins; moist substrate, overuse of the foot pad through increased weight bearing, and reduced activity levels in a captive environment all seem to increase the likelihood of bumblefoot.

Inactivity in captive penguins may lead to increases in bumblefoot

In the wild, Humboldt penguins have been found foraging up to 72km from their rocky nesting beaches in South America (Culik et al. 1998, Chiu et al. 2011) yet in captivity will spend most of their time standing and waiting for food to be brought to them.  Even though zoos meet animal welfare legislation by providing the correct diet and environment for the penguins, in captivity our flightless friends become less inclined to swim unless they are actually fed in the water. Considering their wild cousins spend most of their lives in the ocean, the increased time spent waddling on land is considered just one of the factors to increase bumblefoot onset in these tuxedo wearing birds.

Weird weight distribution of penguins

Bumblefoot is a prevalent condition in non-aquatic species too, with a wide range of research conducted on raptors and the way they spread their weight. The foot of a bird of prey has evolved to stand on a flat surface (think of the wide branches and cliff faces you see in documentaries), but most captive display raptors are tethered to perches, causing them to put pressure directly on their “heel” rather than spreading their weight across the whole foot. Whilst penguins do not even try to stand on perches, their increased time on land has led to some unusual postures. Even before suffering with bumblefoot, captive penguins have been spotted resting back on their heels and raising their toes skywards, as if to take the weight off their toes. This seems to cause a more severe lesion on the main foot pad.

Substrate in zoo enclosures may lead to bumblefoot

bumblefoot in penguins
Wet, hard surfaces? Standing still? The factors leading to bumblefoot in just one photo. Photo by Louise Chiverton

Penguins of all species take to solid ground for two key reasons: breeding and moulting, with some species estimated to spend 75% of their lives at sea (BAS, 2015). Yet in captivity, penguins are found to choose standing over swimming, particularly on flat, unnatural substrates such as concrete or render. In studies on farmed poultry, wet mulchy substrate was linked to higher rates of bumblefoot and when research was undertaken on captive penguins, similar links were made; penguins with wet feet standing on wet substrates had worse feet. Makes perfect sense when you think about it – our own skin cuts easier when wet and infections have easy access to the blood stream. And for penguins, secondary infections can be fatal.

Where do we go from here?

The research all points towards these three factors as the leading causes of bumblefoot. But now, work needs to be done to relieve the pressure off the feet off penguins – both in captivity and in the wild. That’s right, even wild penguins have been spotted with the classic symptoms of bumblefoot and that might be through human influence too. Remember climate change? Well the thought process is that as the ice sheets recede and sea levels start to rise, wild penguins may well be spending more time standing on damp, harsher substrates… just like those in captivity. Whilst further studies need to be completed to properly understand the impact on wild populations, what can be said for certain is that zoo collections must evaluate their enclosure design to reduce bumblefoot onset in the future.

Did you know any of this about penguin health and disease? Let us know what you learned. Also, we more that appreciate any support you can provide so we can continue to provide this type of information to you.

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  1. BAS (2015). Penguins – British Antarctic Survey. [online] Available at: [Accessed 26 Jul. 2019].
  2. Chiu, A.; Cárdenas, S.; Cardeña, M.; Bussalleu, A.; Guerrero, P.; Sandoval, F.; Tremblay, Y., 2011, The route of the Penguin : Use of marine habitat and pattern of attention to the nest by the Penguin of Humboldt ( Spheniscus humboldti ) in Punta San Juan , PeruBoletín Informativo UNOP 6: 21-27.
  3. Culik, B., Luna-Jorquera, G., Oyarzo, H. and Correa, H. (1998), Humboldt penguins monitored via VHF telemetry, Marine Ecology Progress Series, 162, pp.279-286.
  4. Stransky, O., Blum, R., Brown, W., Kruse, D. and Stone, P. (2016). Bumble Foot: A Rare Presentation of a Fusobacterium varium Infection of the Heel Pad in a Healthy Female, The Journal of Foot and Ankle Surgery, 55(5), pp.1087-1090.
  5. Tolpinrud, A., O’Brien, M. F., Justice, W.S.M., Barrows, M., Steele, O. D. M., Gent, S., Meredith, A. ,(2017) Infrared Thermography as a Diagnostic Tool for Pododermatitis in Captive Greater Flamingos (Phoenicopterus roseus), Journal of Zoo and Aquarium Research, 5(1), pp. 48-55.

African Penguins Seriously Affected by Oil Spills

African Penguin

African Penguins Seriously Affected by Oil Spills

by Lydia Alemu

African Penguins are well-adapted for their way of life. With solid bones, flipper-like wings, and webbed feet, they are excellent divers and underwater hunters. Of course, they’d prefer to avoid being eaten in the endeavor to eat, hence their counter shading and well-developed vision above and below the water. They adapt to whatever natural challenges come their way.

Unnatural threats, however, are negatively affecting wild African Penguins. Most of these challenges progress too quickly for penguins to adapt to. As a result, their numbers are decreasing drastically: by 95% over the last century. According to the IUCN Red List, there were 50,000 mature individuals in 2015. In 1910, there were approximately 1.5 million African Penguins. Their numbers continue to dwindle. One major threat is oil spills. African Penguin populations are so fragile that they are only a few disastrous oil spills away from extinction in the wild.

African Penguin

The worst oil spill affecting African Penguins was the MV Treasure

Penguin oil spill

One of the worst South African oil spills to date that affected penguins occurred on June 23, 2000. MV Treasure, a 17-year-old cargo ship, was already damaged by poor weather, old age, and stress when it was towed farther out to sea. Authorities hoped that the move would reduce the impact of an oil spill. Unfortunately, the vessel sank en route with 140,000 tonnes of iron ore and 1,300 tonnes of fuel oil on board between Dassen Island and Robben Island. At the time, these islands respectively housed the largest African Penguin colony of approximately 55,000 individuals and Robben Island has the 3rd largest colony of 18,000 adults.

Volunteers and employees of the South African National Foundation for the Conservation of Coastal Birds (SANCOBB) wasted no time to find, care for, and rehome affected birds. Experienced personnel on oiled bird treatment were flown to South Africa, including staff from the International Bird Rescue in California. Altogether, they relocated around 19,500 unoiled African Penguins to Cape Recife, 800 km away from the oil pollution, with a 98% success rate. (Unfortunately, 241 penguins did not survive the journey.) The cost of this endeavor alone was 300 € ($337.99 USD) per penguin. SANCOBB workers cleaned about 19,000 oiled birds, including adults, and juveniles. They also housed and cared for 3,350 abandoned chicks. 

In total, about 39,000 adult and juvenile African Penguins were handled. This total was over 20% of the total African Penguin population at that time! Within the first six weeks of rescuing African Penguins, volunteers and staff maintained the health of 97% of their charges. Even four years after the Treasure oil spill, 70% of rehabilitated adult penguins, 40% of relocated penguins and 34% of captive-reared African Penguin chicks had returned back to Robben Island! That might even downplay the reality considering that some of the relocated African Penguins might have ended up choosing new islands or beaches to live on since the Treasure sank.

More recent oil spills affecting penguins

Even with the exhaustive effort to help as many African Penguins as possible, staff couldn’t save or even find every penguin affected by the oil spill. An estimated 4,000 chicks as well as 2,000 adult and immature African Penguins died within the first six weeks of the Treasure oil spill. Also, the year’s breeding season was heavily affected with deceased partners, increased time under human care for some penguins, as well as off-kilter molt cycles. Such outcomes would undoubtedly linger for future breeding seasons. 


Imagine another major pollution event on this scale. With their wild population count at approximately 25,000 breeding pairs, their species likely would not survive.


Despite the threat that oil pollution has on African Penguins and other endangered species, two relatively minor oil spills occurred near the South African coastline within the last decade. In 2013, a ship ran aground outside of a nature reserve. About 227 African Penguins were affected. SANCOBB successfully rehabilitated 95% of them.


Most recently, a spokesperson from the Department of Environment, Forestry and Fisheries of South Africa announced that 200 to 400 litres of fuel was spilled on Saturday July 6, 2019. This time, a ship’s fuel tank was over-filled during offshore bunkering. Once again, SANCOBB rushed to help, accepting 21 oiled African Penguins as well as 2 oiled African Penguin eggs within the first three days of the oil spill.

There are things you can do to truly help penguins affected by oil spills

It might seem like there is nothing to be done about oil pollution because the transportation is already in place and regular folks supposedly have little say in or about the industry. However, you always have a say if you actually use your voice. Share your concerns with the governmental leaders in your area to improve protections to regulated nature reserves, for example. When possible, support the creation of marine protected areas rather than new pipelines or other modes of oil transportation. Also, organizations such as SANCOBB are ready to help minimize the detrimental impacts that flaws in current oil transport can have. Donating your time, money, and/or required supplies to them will enable them to do more.

Did you know about these oil spills, the threat to penguins, and the help from volunteers? Let us know.

If you’d like to support penguins, please consider donating to Penguins International so we can help in circumstances like these.

Also read more about penguins in other blogs:


Crawford, R.J.M., Davis, S.A., Harding, R.T., Jackson, L.F., Leshoro, T.M., Meÿer, M.A., Randall, R.M., Underhill, L.G., Upfold, L., Van Dalsen, A.P., Van Der Merwe, E., Whittington, P.A., Williams, A.J., Wolfaardt, A.C. (2000). Initial impact of the Treasure oil spill on seabirds off western South Africa. South African Journal of Marine Science, 22, 157-176.


Barham, P.J., Crawford, R.J.M., Underhill, L.G., Wolfaardt, A.C., Barham, B.J., Dyer, B.M., Leshoro, T.M., Meÿer, M.A., Navarro, R.A., Oschadleus, D., Upfold, L., Whittington, P.A., and Williams, A.J. (2006). Return to Robben Island of African Penguins that were rehabilitated, relocated or reared in captivity following the Treasure oil spill of 2000. Ostrich 77(3&4), 202-209.


BirdLife International 2018. Spheniscus demersus. The IUCN Red List of Threatened Species 2018: e.T22697810A132604504. Downloaded on 14 July 2019.

Zookeepers Help Penguin Conservation

Penguin zookeeper

Zookeepers Help Penguin Conservation

By Megan Spofford

National Zoo Keeper Week 2019 is July 21-27.

Zoos have evolved over the years from being individual organizations that collected animals, to now using their space to house animals for educational purposes or if a species needs human intervention to keep from going extinct. Additionally, zoo workers have developed more of a community and collaborate with each other between facilities, amongst scientists, and with scholars to create conservation programs to save species. This article will delve into some of the work that zoos are doing for penguin conservation.

Penguin zookeeper
Penguin enclosure, London Zoo, Camden, taken 1967
cc-by-sa/2.0 – © Christine

*Warning: Background that is a little dense

Organizations like the International Union for the Conservation of Nature (IUCN) put together teams of experts, many of whom are zoo professionals, to reform policy around the world to benefit wild populations. In the United States, the Association of Zoos and Aquariums (AZA) has developed Taxon Advisory Groups (TAGs) to determine which species deserve the conservation spotlight within their taxa, based on their plight. They then create Species Survival Plans (SSPs) for the chosen ones, and studbooks are used to manage breeding populations who are on SSPs. Studbook keepers are typically working animal caretakers that manage the records in addition to their typical day job. Success stories from these programs have led to captive-bred wild releases to restore populations for black-footed ferrets and Panamanian golden frogs. These are just two examples of how species individuals can be successfully released into the wild through reintroduction programs as long as they are not habituated to humans.

Now the Penguin Conservation stuff!


There is a Penguin TAG that has developed SSPs for ten of the penguin species, which include African, Chinstrap, Gentoo, Humboldt, King, Little, Macaroni, Magellanic, Northern Rockhopper, and Southern Rockhopper Penguins. However, there are currently no reintroduction programs through AZA for penguins. This is because the issues within the environments in which the penguins live need to be addressed before releasing animals into them. Releasing more individuals into the current conditions (oil spills, rising ocean temperatures, food scarcity, etc.) that are causing population declines would not solve the problem. The IUCN has a Species Survival Commission- Penguin Specialist Group that identifies goals to effect change for penguins and their environments through the year 2020 in a report on the IUCN website. The SAFE program developed by AZA has similar goals as the IUCN, but specifically for African Penguins.

“This may even inspire you to take action to help save wild penguins”

penguin zookeeper

Until the time comes for reintroduction programs to be put into action for penguins, TAGs maintain a healthy and genetically diverse population of various penguin species in zoos as “stock.” The penguin “stock” also serve another purpose under human care as ambassadors for their wild counterparts. More simply put: A penguin animal ambassador is a living, breathing example of an extraordinary animal that you would never typically see without traveling to their natural habitat. Getting to see them up close at the zoo brings you into their world, and fosters a connection between you and the animal. Hopefully this causes you to care more about penguins in general. This may even inspire you to take action to help save wild penguins, whether it be by reducing your use of plastics, donating financially to organizations like Penguins International, or environmental tourism to help save a particular species.

If you decide to volunteer with an organization that cares for wild penguin populations, you would likely be working amongst a few zookeepers. At zoos, keepers learn species specific behavior and husbandry, so they are able to transfer those skills while working in situ (Latin word for on site!). An example of this is Maryland Zoo’s work with SANCCOB in South Africa. SANCCOB is an organization that rehabilitates then releases wild seabirds; one of which is the African penguin. During hatching season, which is in the fall, the facility is inundated with abandoned chicks that are reliant on human intervention in order to survive. Zookeepers from Maryland Zoo travel to the facility and contribute their working knowledge to assist in chick-rearing. Smaller facilities that do not have the means to send a keeper to help on site or facilities that are accredited by organizations other than AZA often find conservation groups to fundraise for so that the work can be carried out by field scientists.

More information about penguin conservation projects


You can find out information about conservation projects at specific zoos by looking at their websites or talking with a zookeeper. In honor of National Zookeeper Week (July 21-27, 2019) and the work keepers contribute to animal conservation, we say, “Thanks!”

Did you know all of this about penguins and zoos? Let us know! Also we more that appreciate any support you can provide so we can continue to provide this type of information to you.

Check out some of our other blogs, too, about penguins:


Why Are There No Penguins In The North Pole?

where do penguins live

Why Are There No Penguins In The North Pole?

by James Platt

If you asked a group of people where penguins live, you’d inevitably have a few say that pengiuns live in the North Pole. As we know this isn’t true, but it does beg the question: Why are there no penguins in the North Pole? And for that matter, the entire Northern Hemisphere?

What is the scientific research on why penguins live where they live?

There doesn’t seem to be much scientific research on the matter, so it must be explained using what is known about penguins around the world and then given the best answer possible. There are 18 species of penguins, of which 7 of them live in the Antarctic such as the Emperor Penguin (Aptenodytes forsteri) and King Penguins (Aptenodytes Pategonicus). The Yellow-eyed Penguin (Megadyptes antipodes) and Royal Penguin (Eudyptes schlegeli) are 2 of 6 species in Australia and New Zealand. There is one African Penguin (Spheniscus demursus) and 3 species that inhabit the Americas with the Galapagos Penguins (Sphendiscus mendiculus) living just slightly in the Northern Hemisphere, living so close to the equator.

Penguins live (almost) entirely in the Southern Hemisphere

With so much variety among penguin species it seems strange that they never moved farther north and filled more ecological niches as they did in the Southern Hemisphere. They found a home on the Australian and African continents where temperatures can be as hot as anywhere on earth, so temperature isn’t as much of an issue as most people may think. King or Emperor Penguins may not find the heat as easy to deal with as they are still adapted for the harsh environment of the Antarctic. However, the Galapagos Penguins, Little Penguins (Eudyptula minor) and African Penguins can withstand much hotter temperatures and could theoretically make the shift farther north.

Galapagos penguins
Figure 2: Volt Collection/Shutterstock

Looking at the Galapagos Islands, they are surrounded by hundreds of miles of open ocean. The larger varieties could swim that far if they were searching the Southern Ocean for food, but the smaller Galapagos Penguin doesn’t have a chance of swimming to Costa Rica or El Salvador which is why they have remained isolated to the islands (Heath and Randall, 1981). The same goes for the Snares Penguin (Eudyptes robustus) and Fiorland Penguin (Eudyptes pachyrhyncus) along with all the other species in Oceania if they were to move up through Papua New Guinea and Indonesia.

Some species may be adapted to the heat, but they are not adapted to move fast on land, so they will only stick to the coast and won’t often move north unless they are forced to. A behavioural study investigated how the African Penguin deals with heat and they spend most of their day in the ocean and only return to land in the late afternoon (Frost et al, 1976). At night, they are much less likely to move farther north in search of territory as it is more dangerous at that time. In South Africa and Namibia, the African Penguin also has a lot more predators to be wary of, making any move farther north a risky one. In the Antarctic, when the penguins are on the ice they have no real threats from predators, so they have not evolved much protection against land predators. But on these other continents they can be real threats, especially to the chicks.

In Antarctica, penguins and their nest sites are left relatively untouched by humans and they are left to breed and live in peace. African Penguins have been pushed to the brink by humans taking their guano for fertiliser and trampling their burrows (Trathan et al, 2014). Many of the southern island species were hunted for oil until the last century. So, life outside of the Southern Ocean isn’t great for many of the species. It might be that penguins couldn’t survive any more outside pressure from humans and any populations that have successfully moved farther north were pushed to extinction before they could gain a proper foothold in the area.

All these reasons combine to make a compelling case for why there are no penguins in the Northern Hemisphere. It’s likely since penguins won’t move over large distances on land, they struggle to find new territory farther north. Any that do make it are likely either scared off by curious humans or killed by predators as they will make an easy meal for many land predators. The Galapagos Penguins probably won’t make it any farther due to their geographic location being so far from anything else. If they were on the other side of South America then they very well could have used their adaptation to hot climates and island hopped all the way to North America through the Caribbean and Cuba.  The closest thing to a penguin that did live in the North Pole was the Great Auk (Pinguinus impennis) and was subsequently hunted to extinction in the 1800s. Just as the Dodo (Raphus cucullatus) had no fear of humans because they hadn’t ever faced predators, the Great Auk suffered the same fate. So maybe its better if penguins stay South and remote for now.

Royal Penguins
Figure 3: Sonja Ross

What do you think? Is there another reason they haven’t moved farther north? Leave us a comment. Also, please help us continue to learn more about penguins by donating to Penguins International.

Check out some of our other blogs, too:

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Frost, P., Siegfried, W. and Burger, A. (2009). Behavioural adaptations of the Jackass penguin, Spheniscus demersus to a hot, arid environment. Journal of Zoology, 179(2), pp.165-187.

HEATH, R. and RANDALL, R. (1989). Foraging ranges and movements of jackass penguins (Spheniscus demersus) established through radio telemetry. Journal of Zoology, 217(3), pp.367-379.

Trathan, P., García-Borboroglu, P., Boersma, D., Bost, C., Crawford, R., Crossin, G., Cuthbert, R., Dann, P., Davis, L., De La Puente, S., Ellenberg, U., Lynch, H., Mattern, T., Pütz, K., Seddon, P., Trivelpiece, W. and Wienecke, B. (2014). Pollution, habitat loss, fishing, and climate change as critical threats to penguins. Conservation Biology, 29(1), pp.31-41.

Do Penguins Have Knees?…and other frequently asked questions

Penguin knees

Do Penguins Have Knees?…and other frequently asked questions

by Autumn L. Syracuse, Educator I

“Do penguins have knees?” and “So what is a penguin?” are two of the most common questions I hear regarding our penguin colony on display at the Aquarium of Niagara ( Before we talk about penguin anatomy, let’s discuss what a penguin is exactly, first. It’s hard to imagine that these chunky bipeds that don’t fly — but swim — are indeed birds. Ostriches, emus, and rheas are pretty easy to identify as birds with their fluffy plumage of feathers. So why does it seem odd to include penguins in this group of flightless birds? Are all birds descended from one common ancestor?

Birds are: warm-blooded, air breathing, egg-laying, covered in feathers, and possess a bill. Now let’s take a look at penguins: They check all the boxes! “But why are they birds? They are so funny looking!” Although outwardly different from most other birds, penguins still possess many characteristics of other avian species. Let’s get down to the bones of it.

Penguin knees

Okay, so do penguins have knees?

A penguin’s skeletal structure is laid out in the same general pattern as other birds. One obvious characteristic of a bird skeleton is the keel, or sternum. This is designed to be very wide and flat, but lays perpendicular to the ribs. This large bone helps to attach the flight muscles and tendons, which is very important in both form and function to flight. And since penguins “fly” through the water, which is denser, they too need to rely on the keel and flight muscles for propulsion. 

Another important adaptation that varies from other flighted (volant) birds is the density of their bones. Most birds we see flying in our yards and neighborhoods have skeletons with bones that are hollow. This creates a skeleton that is extremely lightweight, allowing the birds to be able to lift off into flight. Penguins would not benefit from bones of this same density. Penguins need to be able to dive underwater to hunt for their fish, and hollow bones would make them too buoyant. To help with this, penguin bones are solid and heavy, helping to give them more weight in order to dive deep.

Penguin knees are tucked up inside their body

Penguins are designed to be streamlined and hydrodynamic, so having long legs would add extra drag. Having short legs with webbed feet to act like rudders, helps to give them that torpedo-like figure. If we compare bird anatomy with humans, we would see something a bit peculiar. By taking a look at the side-by-side image in Figure 1, you can see how their leg bones compare to ours. What most people mistake for knees are actually the ankles of the birds. This gives the illusion that bird knees bend opposite of ours. The knees are actually tucked up inside the body cavity of the bird! So how does this look inside of a penguin? In the images below, you can see boxes surrounding the penguins’ knees. 

Penguin knees
Penguin knees
Colored boxes highlighting the location of penguin knees.

Imagine yourself wearing an oversized shirt, and pulling it over your knees so that only your ankles and feet are showing. Now imagine you’re trying to walk forward in this position. I bet you’d waddle too! This design gives the penguin an advantage in the water to help them swim quickly to catch food or avoid predators. On land, they tend to be slower and clumsy, which makes them more prone to predators. For Antarctic penguins, they rarely encounter predators on land, so having larger bodies isn’t detrimental. For other species in temperate or tropical climates, the water tends to be a bit safer place.

Originally, penguins were classified in the same group as other flightless birds (Ratites). After multiple studies, it was discovered that penguins evolved from flying birds, which were separate from the ancestors of other flightless birds. Mitochondrial DNA has further suggested their relationship to other seafaring flighted birds such as albatross, frigatebirds, and loons. In 2006, more genomic testing suggested that birds of the Ciconiiformes order (storks, gannets, plovers, and boobies) were their closest living relatives (Watanabe et al. 2006). 

Paleontologist Ewan Fordyce with a model replica of a Waimanu penguin.

It is still uncertain as to what other bird species may be related to penguins. But from finding and examining their fossilized bones, we have an idea of what they were like nearly 60 million years ago. Known by the genus name Waimanu, these ancient penguins may hold the record for the oldest evidence of bird lineage. Scientists believe that the extinction event that wiped out the dinosaurs during the Cretaceous period, also eliminated almost all bird species. After this catastrophe, it is believed that modern day penguins evolved from the few species that had survived, evolving quickly over a short amount of time—in relation to Earth’s history (Fordyce and Ksepka, 2012).

“So what kind of animal are they?” All of the evidence points to birds, but it still leaves questions unanswered. After learning of this fossil evidence, it leaves me with this question: “Are penguins birds? Or are birds penguins?” When we take a look at other body systems and explore their behavior, things become more “black and white.”

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Have you ever wondered before reading this if penguins have knees? Let us know what you think, and what you learned. Also, please help us continue to learn more about penguins by donating to Penguins International. We more than appreciate your support!

You can also read more about penguins in the following blogs:

Flipper Bands – Are they worth the risk to save a species?

penguin flipper bands

Flipper Bands – Are they worth the risk to save a species?

by Georgia Podmore

You can walk around a variety of zoos and see that many penguins will have flipper bands with the purpose of enabling keepers to identify the individuals easily. At a glance, this might not appear to be a problem. The penguins are swimming in their enclosure and seem to be behaving normally. However, the small metal band that has an identity scribed onto it could be causing more harm than it is worth.

Since the 1970s, research has investigated the use of flipper bands and the harm that they could cause, generally focusing on wounds that are caused when a penguin goes through its annual molt (Hampton, Ryan and Underhill, 2009). It is not only penguins in captive environment that may be having trouble with flipper bands. Researchers are continuously trying to uncover new information on different species, population counts and climate change. To gain significant data, flipper bands are often used when studying large colonies of penguins.

Are Flipper Bands Safe?

For many years, this topic has caused a debate among researchers. The team at the University of Strasbourg used evidence to back up Rory Wilson’s statement that “Some tags seem to slow down penguins”. (Jackson and Wilson, 2002) The researchers placed metal bands on 50 King Penguins, while 50 other penguins were fitted with minor radio-frequency transponders. After 10 years, it was revealed that survival rates for the banded penguins was 16% lower than the penguins that were fitted with transponders. It was observed that banded birds were affected in a variety of ways, such as being slower to breed and taking longer periods of time to forage (Gauthier-Clerc et al., 2004).

A Yellow-eyed Penguin with a flipper band for identification.

Although there are a variety of studies examining flipper bands and the effect on wild birds, some studies have found no evidence to back this up. Jackson and Wilson (2002) found that with Royal Penguins, there was no difference in the growth of chicks, survival in harsh climates or success of breeding in flipper-banded birds compared to transponder-fitted birds. However, the main portion of research – that has concluded that flipper bands are not detrimental to penguins – has been evaluating short-term use of them (Saraux et al ,. 2011).

Dee Boersma, a leading penguin researcher, believes: “All bands and all penguins are not the same” (Culik, Wilson and Bannasch, 1993).  It is hard to settle a debate when research looking into the effect of flipper bands on penguins is focusing on a variety of different species in a diversity range of environments. With 18 species of penguins all living in various climates, it is difficult to come to a concluding factor on the effect of flipper bands on welfare, as it is dependent on the individual penguins, as well as what type of flipper band is used.

Flipper bands are currently used in research to gain knowledge on the effects of climate change and the impact it is having on marine mammals. Scientists need to gather information on climate change, as it is believed that through observing marine mammals, predictions can be made earlier. Climate change is one of the biggest threats to the world and the species living on it. If scientists can make predictions earlier, this can hopefully make a positive change. If they are unable to use the flipper bands to research the penguins, many other species may be in danger of becoming extinct. Dee Boersma states: “we do have to do some harm if we want to follow individuals” (Culik, Wilson and Bannasch, 1993). However, is it worth the risk to save a species? Significant data shows that penguin survival and breeding rates become lower when fitted with a flipper band. Could the long-term studies of climate change cause the extinction of penguins if these types of research are continuously used?

A Northern Rockhopper Penguin with a flipper band

Investigations into the use of small transponders have found that these have a less negative effect on the individual penguins, although further research is needed to understand whether this will affect welfare negatively in the long-term. The topic is debatable in many ways as studies have found different statistics. It is obvious that flipper bands do cause harm – whether minor or major to penguins – but climate change research is necessary to save the penguins. A study that may help push the use of transponders or similar tracking devices may be a project that assesses captive penguins. The penguins would all be the same species and live in the same environment. This research would enable more data to be published on the effects of flipper bands on specific species, climates and other factors such as age and gender. Through this type of research, this may then enable scientists to lower the risk of affecting wild penguins if they are able to understand what factors may increase the risk.

What are your thoughts on the use of flipper bands on penguins? Let us know! Also, please help us continue to learn more about penguins by donating to Penguins International. We more than appreciate your support!

You can also read more about penguins in the following blogs:


Culik, B., Wilson, R. and Bannasch, R. (1993) Flipper-bands on penguins: what is the cost of a life-long commitment? Marine Ecology Progress Series, 98, pp.209-214.

Gauthier-Clerc, M., Gendner, J., Ribic, C., Fraser, W., Woehler,E., Descamps, S., Gilly, C., Le Bohec, C. and Le Maho, Y. (2004). Long-term effects of flipper bands on penguins. Proceeding of the Royal Society of London. Series B: Biological Sciences, 271 (supple_6)

Hampton, S., Ryan, P. and Underhill (2009) The effect of flipper banding on the breeding success of African Penguins Spheniscus demersus at Boulders Beach, South Africa. Ostrich, 80(2) pp.77-80.

Jackson, S. and Wilson, R. (2002) The potential costs of flipper-bands to penguins. Functional Ecology, 16(1), pp. 141-148.

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