March of the Fossil Penguins

Fossil penguin discoveries and research

Penguins are not fans of opera?

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Written by Dan Ksepka

May 15, 2016 at 7:23 pm

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Reading the Mind of the Oldest Penguin

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The fossil skull (left) and a virtual endocast (right). Figure from Proffitt et al. (2016).

Today is World Penguin Day, which makes a good excuse to catch up on fossil penguin blogging. Recently, James Proffitt and colleagues published a study on fossil penguin neuroanatomy. As we’ve discussed previously, the brain morphology of extinct animals can be reconstructed by computed tomography (CT) scanning, which allows researchers to map out the volume and shape of the brain based on the cavity it once occupied in the skull. This paper provides the first look at the neuroanatomy of the early penguin Waimanu.
Waimanu is the oldest reported penguin taxon. It is known from Paleocene rocks in New Zealand, where two species have been discovered (Waimanu manneringi and Waimanu tuatahi). The skull that was scanned for this study belongs to the genus Waimanu, but whether it represents one of the two known species or a third new species has not yet been untangled.  Even though Waimanu is a very early penguin, probably representing a point in the evolutionary history of penguins just a few million years after the loss of aerial flight, it already shows many of the features that are typical for penguin brains. These include the widening of the cerebellum, and the lack of any impressions of cerebellar folds on the endocast (interestingly, modern penguin brains do have cerebellar folds like other bird brains, and it has been suggested that the lack of folds on the endocast is due to the “cushioning” of the brain by meningeal tissue).  On the other hand, Waimanu is primitive in showing weak development of the “Wulst”, a structure associated with complex visual processing and other functions. The Wulst becomes expanded later in penguin evolution, as seen in some of the endocasts I have studied from Eocene and Miocene penguins.

An evolutionary tree mapping endocast shape in aquatic birds.  Figure from Proffitt et al. (2016)

These new data let us push deeper into the history of penguin brain evolution. As we get closer and closer to the loss of aerial flight by probing deeper down the trunk of the tree, it will be interesting to see when some of the “penguiny” brain features that are already present in Waimanu first arose.  As a whole, endocast studies of penguins are a great example of how new research becomes possible with new technology. Five years ago, there were no fossil penguin endocasts at all, and today we have five and there are plenty of additional fossils that have already been scanned or are waiting patiently for their turn in the CT lab. Perhaps by the next World Penguin Day we will have ten endocasts?
Proffitt, J. V., J. A. Clarke, and R. P. Scofield. “Novel insights into early neuroanatomical evolution in penguins from the oldest described penguin brain endocast.” Journal of Anatomy (2016).

Written by Dan Ksepka

April 25, 2016 at 10:12 am

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(Sub)fossils in Guano

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Beneski_Amherst 010.jpg

This week, I visited the Beneski Museum of Natural History at Amherst College. The museum is home to one of the most famous fossils in the world, a slab bearing the first fossil dinosaur footprints ever discovered. Remarkably, the slab was used for years as a doorstop before its importance was recognized.

While the museum’s remarkable assemblage of fossil trackways is the most unique aspect of the collections, a different type of fossil caught my eye. Within a nifty display illustrating different pathways to fossilization, a somewhat unfortunate-looking bird sits in a pull-out drawer. The specimen was collected from Guanape Island, off the coast of mainland Peru. Although it looks almost like a mangled rubber chicken, the specimen is actually hardened and pretty much reduced to the bones. The bird is encased in a thin layer of guano: that is, it is literally preserved in poop. Guano is the term scientists use for the bird and bat droppings. Guano is rich in phosphate, which under the right conditions can enhance preservation of shell and bone. In places where animals congregate in large numbers, guano may form layers than are meters and meters deep.

Guano Penguin.jpg

A closer look reveals the specimen here is a penguin: the dead giveaway is the very short wing.  This penguin is most likely a Humboldt Penguin, a species which inhabits the coasts of Peru today and happens to feel quite comfortable digging into guano layers to create burrows for nesting. Guano nesting can be hazardous, however. Humans have recognized that guano makes an excellent fertilizer for centuries, and have often excavated guano deposits with little regard for the birds using the deposits as breeding colonies, stripping them bare with rapacious greed. This approach is short-sighted, as the deposits are replenished by the birds themselves – more responsible harvesting schedules can turn guano into a renewable resource and have been adapted in recent times and many sights.

Written by Dan Ksepka

March 8, 2016 at 11:58 am

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How many feathers does a penguin have?

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How many feathers does a penguin have? It seems like a straightforward thing to address, but it is also exactly the type of basic natural history question that often gets overlooked. It appears that no one had taken the time to thoroughly calculate the number before. Surprisingly, previous accounts ranged from 11 feathers per square centimeter to 46 feathers per square centimeter, and did not even agree on what types of feathers the penguins have.


A (delightfully named) feathergram showing feathers from different parts of an Emperor Penguin. Image from Williams et al. (2015).

Feathers fulfill many functions beyond flight, including thermoregulation, camouflage, display, and waterproofing. Different types of feathers contribute to these functions. Contour feathers are large and stiff-vaned feathers that generally form the outer layer of a bird’s feather coat. Flight feathers are the contour feathers than create the airfoil of the wing. These feathers are large in volant birds, but in penguins the “flight” feathers are reduced to tiny, scale-like structures. Plumules are downy feathers with looser barbs and a soft, cottony texture. These feathers typically lie under the contour feathers and provide an insulating layer. Perhaps the weirdest type of feather is the obscure filoplume. Filoplumes are essentially small, bare feather shafts. They can serve no aerodynamic or insulatory purpose and are instead hypothesized to help birds detect the orientation of their larger feathers. Penguins have long been considered to lack filoplumes, but that turns out not to be the case.


A tiny filoplume, nestled at the base of a contour feather. Image from Williams et al. (2015).

A new study tackled the question of Emperor Penguin feather density, a compelling issue given the ability of these birds to survive extremely cold and windy conditions. The team salvaged Emperor Penguin carcasses from Antarctica and took small square patches of skin from different parts of the body.  Within each patch, they counted and mapped out the arrangement of different feather types. Whereas previous scientists had overlooked filoplumes in penguins, the team observed them nestled at the base of larger feathers. In fact, it seems that each contour feather on the body has its own associated filoplume. Overall, the most numerous feathers were the downy plumules, which were four times more abundant than contour feathers and filoplumes. This surfeit of downy insulation makes sense given the harsh environment Emperor Penguins inhabit. It may also play a role in locomotion, as a place to store air to release when speeding towards the surface (see more about Bubbling Penguins here).

So how many feathers does an Emperor Penguin have? It turns out density varies quite a bit around the penguin, from as low 5.8 feathers per square centimeter on the back of one penguin to 13.5 feathers per square centimeter on the front of another. The researchers extrapolated from their samples that the full body would have 144,000 to 180,000 total feathers.  Any volunteers to count them individually and confirm?


Reference: Williams CL, Hagelin JC, Kooyman GL. 2015 Hidden keys to survival: the type, density, pattern and functional role of emperor penguin body feathers. Proceedings of the Royal Society B 282: 20152033.

Written by Dan Ksepka

February 2, 2016 at 11:48 am

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The Penguin’s Palette

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I’ve written an article on color in penguins, highlighting some of the remarkable discoveries my colleagues in the modern penguin world have made about pigments, structural color, and more (plus of course a mention of fossil penguin feathers). The article is in the January/February 2016 issue, and you can check it out here as well:

Written by Dan Ksepka

January 24, 2016 at 12:00 pm

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Ultraviolet Beaks

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Today is Penguin Awareness Day. Let’s discuss a feature of which we humans may not be aware, because of our limited visual perception. Our eyes can detect the visible light part of the electromagnetic spectrum, spanning the range from about 700 nanometers (red) to 400 nanometers (violet) wavelength.  Many birds, including penguins, see beyond this range into the ultraviolet portion of the spectrum.  Birds often have “hidden” markings that they themselves can see, but can only be detected by humans through artificial illumination.


King Penguin, Falkland Islands. Photo by Ben Tubby.

King Penguins are one species that have ultraviolet markings, as scientists have discovered. These large penguins are already stunning birds, with orange patches of color along their necks, ear regions, and the sides of their beaks. Recently, scientists delved deeper to detect ultraviolet patches are also positioned along the lower bill. Both the visible and ultraviolet colors appear to play a role in attracting mates.

How do King Penguins produce ultraviolet colors? The answer is multilayered reflector photonic microstructure. Essentially, the outer layer of the beak contains alternating layers of high refractive index and low refractive index materials. Reflected light from the different layers interacts to bounce back wavelengths in the ultraviolet spectrum. Research by Dr. Birgitta Dresp-Langley and colleagues has revealed that King Penguin beaks have a layer filled with special folded microstructures and intervening filaments
of β-keratin. These markings help indicate maturity, and may also be attractive to other penguins.  As a King Penguin grows, the ultraviolet hue of the beak markings increases. Surveys of wild penguins  show they are strongest in recently formed male–female pairs. When scientists hid the ultraviolet markings by painting a layer of varnish over a penguin’s beak, those birds had a harder time finding a mate – perhaps the equivalent of the penguin hitting the local watering holes without enough lipstick or cologne!


King Penguin, Falkland Islands. Photo by Ben Tubby.


Dresp, B., P. Jouventin, and K. Langley. 2005. Ultraviolet reflecting photonic microstructures in the King Penguin beak. Biology Letters 1: 310–313.

Dresp, B, and K. Langley. 2006. Fine structural dependence of ultraviolet reflections in the King Penguin beak horn/  The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology 288: 213-222.
Jouventin P., P.M. Nolan, E.S. Dobson and M. Nicolaus. 2008. Coloured patches inXuence pairing in king penguins. Ibis 150:193–196

Written by Dan Ksepka

January 20, 2016 at 5:50 pm

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Penguins Taste Poorly

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A recent study suggests penguins may have a poorly developed sense of taste. This is not to be confused with tasting bad – which accounts by sailors forced to survive on penguin meat suggest is also true.  Rather, new research suggests penguins lack the ability to detect sweet or savory tastes.

Humans perceive 5 basic categories of taste: sweet, sour, salty, bitter, and umami (which might best be described as savory). However, evidence suggests penguins are limited to just sour and salty. Researchers came to this conclusion after combing through recently assembled genomes from two penguin species. A mutation in the umami taste receptor gene Tas1r1 causes it not to function properly in penguins. Tasr2r genes, responsible for bitter reception, are non-functional in penguins too. Penguins also lack the taste receptor gene Tas1r2, which is associated with sweetness detection. These factors would preclude them from enjoying, say, a Shake Shack burger or a candy apple.  Penguins are not alone in lacking the ability to taste sweet – Tas1r2 appears to be absent in all birds, though nectar-loving hummingbirds seem to have repurposed Tas1r1 to detect sweetness.


This Gentoo Penguin chick will never know the joy of a Snickers Bar. Photo shared via Wikipedia by russavia.


Are penguins hindered by their diminished sense of taste? Probably not. As the authors note, penguins swallow their food whole, so they might not benefit too much from telling whether or not it tastes good. Truth be told though, I have certainly known a few aquarium penguins who strongly prefer one type of fish to another when it comes to their dinner – so much so that their keepers serve the less desirable portion first while they are still hungry.

There is one aspect of the study that clashes with paleontological evidence. The authors conclude the loss of some tastes may be related to the origin of penguins in Antarctica. However, when one considers the complete penguin family tree including the many extinct species, it is far more likely penguins originated at lower latitudes when climate was much warmer and invaded icy Antarctica late in their history.

One exciting extension of the research is that we now have a reason to test the conclusions in the real world. Perhaps some biologists are now hatching a plan to tempt penguins with sour pickles, salty chips, or sweet pieces of candy to see if they can indeed tell the difference.

Reference: Zhao, H., J. Li, and J Zhang. 2015. Molecular evidence for the loss of three basic tastes in penguins. Current Biology 25 :R141 – R142

Written by Dan Ksepka

December 3, 2015 at 4:21 pm

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