From Will’s Skull Page, here.

Here’s a skull of a wild boar. Note the loooong face, practically a straight line from the tip of the snout to the top of the back of the head.

We shall now proceed through a series of pig skulls with increasingly steep foreheads.

From the UCL Museums and Collections blog, here.

Some domestic pigs have a longish snout and nearly straight forehead, like their wild forebears. (Or foreboars, if you will.)

A cast skull from Carolina, available here.

But it seems–from a quick, unscientific, and in-no-way-standardized image search–that the vast majority of domestic pigs have at minimum a more steeply-inclined forehead.

This one was auctioned in New Zealand, at this site.

Foreheadization is becoming undeniable.

From skullbase.info, here.

Is this one any more pronounced than the one before? I’m not sure, and so far I’m too lazy to try superimposing the skulls. But they don’t even look like the same kind of animal as the wild boar shown at top.

From theweirdandwonderful.com, now apparently only available on Pinterest, here.

In my explorations so far, this appears to the ne plus ultra of short-faced, high-forehead domestic pigs, excluding truly pathological cases. The line from the inflection point of the forehead to the occiput is twice the length of the snout!

From theweirdandwonderful.com, now apparently only available on Pinterest, here.

Oddly enough, the high forehead in domestic pigs is not always associated with a super-short snout, as this skull demonstrates.

This figure from Owen et al. (2014) sums up the shape differences between domestic (left) and wild (right) Sus scrofa.

Okay, so domestic pigs have shorter snouts and steeper foreheads than wild pigs of the same species. But y tho? It seems to be part of the “domestication syndrome” present in many domesticated animals, which includes a shortened snout, smaller teeth, piebald coloration, floppy ears, a curly tail, and a host of other morphological and behavioral traits. Interestingly, pigs seem to show more aspects of domestication syndrome than any other domestic animals other than dogs, as shown in the figure below, from Sanchez-Villagra et al. (2016).

Okay, so domestication, but how? It’s not like the Domestication Fairy comes in the night and steals half your snout.

Wilkins et al. (2014: fig. 1)

The various morphological changes that go along with domestication syndrome seemed disconnected until 2014, when Wilkins et al. proposed a pretty nifty hypothesis, which goes like this:

  • Probably the most crucial aspect of domestication is selection for tameness, which is really selection for reduced adrenal gland and sympathetic nervous system activity, so the animals aren’t freaking out all the time.
  • The adrenal glands and sympathetic ganglia are derived from embryonic neural crest, which also influences the growth of the teeth, brain, skull, vertebral column, and ear cartilages, and the distribution of melanocytes in the skin and coat.
  • Selection for increased tameness (= reduced freaking out) is really selection for reduced neural crest activity in early development, and the smaller teeth, shorter snout, floppy ears, curly tail, patchy coloration, and so on, are unselected developmental consequences of reduced neural crest activity.

Wilkins et al. (2014: fig. 2)

So far, so good. The neural crest hypothesis seems to have genuine explanatory power, in that it lassos a disparate set of phenomena and provides a single, logical cause. Of course not everyone is convinced, and the neural crest hypothesis could be true without ruling out other complementary mechanisms and confounding effects. Along those lines, Sanchez-Villagra et al. (2016) is worth a read. It’s free at the link below, as is Wilkins et al. (2014).

The neural crest hypothesis might explain why domestic pigs have shorter snouts than their wild relatives, but I think there must be some other factors in play to explain pig foreheads. Which is fine, domestic dogs have a staggering variety of skull shapes that reflect thousands of years of strong artificial selection, and probably a healthy dose of unintended consequences and other knock-on effects. Given that pigs have been domesticated for a long time, were probably domesticated many times in many places, have had frequent infusions of wild-type genes (from possibly genetically disparate wild populations), and have been canalized into different breeds, it might actually be weirder if they all looked like short-snouted wild boars. All of which is a long way of saying that I’m not surprised that domestic pigs don’t all fall on some morphogenetic monocline from wild boars, but I’m still curious about how they got their foreheads.

I actually started writing this post before the very interesting discussion of pig domestication flared up in the comments on Mike’s pig skull post. Mike’s two skulls nicely illustrate the difference between forehead-less and, er, forehead-ful conditions, and the comment thread touched on a lot of related issues and is worth a read. In particular, I’d like to note again that domestic pig skulls are not notably paedomorphic with respect to wild boars, other than having short snouts–they’re on a different morphogenetic trajectory (Evin et al. 2016).

For a nice comparison of domestic pig and wild boar skulls, see Marcus Bühler’s post at Bestiarium, here.

References

Long-term readers will remember that way back in the pre-history of this blog, I wrote about my experience de-fleshing a pig head, which because the very first part in our ongoing series Things to Make and Do. In a subsequent post with a sheep-skull multiview, I included the multiview of that pig skull, too. Here it is:

Mike’s first pig skull, cranium only. Top row: dorsal view, anterior to right; middle row, from left to right: posterior, right lateral and anterior views; bottom row: ventral view, anterior to right.

As I noted in that sheep-skull post, I no longer own that skull: I donated it to be the first prize for the quiz in the very first TetZooCon, and it was won by Kelvin Britton.

But around the same time, our church hosted a barbecue even in which an entire pig was slow-roasted, and at the end of it I took the head home and prepped the skull out of it. The bone was much more fragile for having been roasted instead of simmered, and was in some danger of crumbling apart, but I stablised it with diluted PVA and it holds together OK.

Here it is:

Mike’s second pig skull, cranium and mandible in articulation. Top row: dorsal view, anterior to right; middle row, from left to right: posterior, left lateral (reversed) and anterior views; bottom row: ventral view, anterior to right.

Even allowing that the new skull was photographed with the mandible in place, the difference between the two is shocking. In particular, check out the dorsal views: the zygomatic arches of the first pig protrude way further laterally, and are much more robust than those of the second pig, and the whole shape of the skull roof is different.

I’m not sure what to make of this. I assume what we’re seeing here is variation of different breeds within the single domesticated species Sus domesticus, analogous to the way bulldog and greyhound skulls differer dramatically despite both being breeds of Canis familiaris. There are a lot of pig breeds out there, so perhaps it’s not too surprising. On the other hand, while the different dogs were bred for different purposes, I’d have thought all the pig were bred for the same purpose: to put on weight and provide meat. So I don’t know why such different skulls would have been selected for.

Well, it’s time. Ten years and almost 5 months after Mike kicked off our “Things to Make and Do” section with his post on cleaning a pig skull, I am finally getting around to prepping a pig skull of my own. There will be a complete play-by-play coming, but for now I want to focus on what is usually the least-pleasant step in prepping a skull: extracting the brain. Aside from the relatively small and often tortuous passages for the cranial nerves, the braincase is a cul-de-sac, with a big glob of tissue (the brain and associated meninges and vessels) only accessible through a relatively small hole at the back of the head (the foramen magnum). Virtually every tutorial and how-to on prepping skulls has some section where the author advises you to basically swirl something around in there, get stuff out the best you can, and prepare to deal with a lot of nastiness along the way. So I had my antennae out for anything that might help, and in the local dollar store I ran across the beauty shown above.

I figured drain rooter = brain rooter, and I was only risking a buck, so I picked one up. It worked a trick: by putting the pig’s snout down the drain, running hot water into the foramen magnum to continually flush out the loose bits, and vigorously exploring the cranial cavity with the brain rooter, I was able to get the whole brain out in about 10 minutes. To be clear, all the tissue came out the foramen magnum; there would be no way to get it to come out the nose without breaking the ethmoid bone and destroying the nasal turbinates. I only put the head snout-down for ease of access. I had a great deal of control, and I could tell pretty well which areas were getting emptied out and which still needed work. All I missed was a small glob of meninges and dural venous sinuses, which came out easily after the first simmer.

Some specific advantages of the drain rooter as a brain extractor:

  • backward-pointing teeth to hook out the tissue
  • flexible plastic so you can go pretty hard with it without damaging the bone
  • super long so you’re not going to find a job too big for it, OR you can cut it to length
  • still works for unclogging drains
  • dishwasher safe
  • dirt cheap

Go have fun.

UPDATE: Turns out pigs have an insane amount of cartilage and mucosa in their nasal cavities, and the brain rooter is pretty good at getting that stuff out, too.

The stupidest head

August 21, 2019

Left: Homo sapiens, head, neck and upper trunk in right lateral view (unprepared specimen). Right: Camarasaurus sp., skull in left lateral view. Photograph at the Natural History Museum of Utah, Salt Lake City, Utah. 2016.

Last September I spent a day in the LACM Herpetology collections with Jessie Atterholt, looking at weird features in crocs, lizards, snakes, and salamanders. I’ll have more to say about the specific things we were looking for in a month or so, after Jessie’s given her talk at SVPCA. This was just an incidental hit. We were looking at cryptobranchid (literally “hidden gill”) salamanders, because they’re big enough that you don’t need a microscope to see all their weird features. Cryptobranchids include the North American hellbender, Cryptobranchus alleganiensis, and the giant salamanders of China and Japan, Andrias davidianus and Andrias japonicus, respectively, plus a raft of fossil taxa.

This is the mandible of LACM 162475, a specimen of Andrias davidianus, in right lateral view. I’d never spent quality time with the skeleton of a giant salamander, and I was impressed with how evil their teeth are. Just in terms of general outline, these little murder-sticks wouldn’t look out of place in the jaw of a dromaeosaur. Click to enfangenate.

Jessie did an Instagram post on the upper jaw of Cryptobranchus a few months ago, and as long as you’re over there, have a look at the half a pig head that she just plastinated for our colleagues in WesternU’s College of Veterinary Medicine. It’s not the same pig as the one we hemisected last December, but I think it got cut at the same time.

My friend and frequent collaborator Jessie Atterholt has her office in the next building over from mine. When you walk in, you see something that looks approximately like this. Not exactly like this, because I took these photos in February and she’s changed a few things (and I’m rubbish about getting stuff posted in a timely fashion).

The last time I showed an office full of amazing stuff like this, it was Peter Dodson’s. It will come as no surprise that Jessie was Peter’s student at UPenn before she went to Berkeley for her PhD.

The far case holds mostly books and skulls. Dr. A has her own plastination setup for making preserved organs and organisms, and the snake on the second shelf here is one that she prepped herself. One side of the snake still has the skin on, the other half has been skinned to show the muscles. This is crunch week for me so I don’t have time to ID all of the stuff, but alert readers should have no problem spotting some digitally-resurrected Haplocanthosaurus bits.

Mostly skulls on the middle rack. The sirenian skull on the second shelf and the cave bear on the fourth are both casts, but almost everything else is real bone. The bighorn sheep on the middle shelf is a natural mummy.

Here’s a close-up of the top shelf. Other than some 3D-printed human skull bones sitting in front of the brain slice on the left, everything here is real bone, including the lion, baboon, and human skulls, and the giraffe cervicals winding across the top. Jessie’s been collecting since she was a kid and the African megafauna are gifts from a globe-trotting family friend.

The upper shelves here have quite a few of Jessie’s plastinated specimens, both whole organisms and things like hearts and kidneys from various critters.

A close-up of some of Jessie’s coolest anatomical preparations. In back is an internal cast of the lungs and bronchial tree of a cat. The baby rattlesnake died after eating a proportionally gigantic lizard — I was dumb and forgot to flip the snake over to show the lizard inside, plastinated along with its predator. The ground squirrel on the right is another half-fleshed, half-skinned plastinate, and the mouse up front is a classic dissection presentation, preserved forever through plastination.

I’ve heard it said that the difference between a collector and a hoarder is curation. As someone who definitely lurks more on the hoarder end of that spectrum (to paraphrase Dave Barry, if you could see my office you’d be blinded or driven insane), I’m pretty darned jealous of both the breadth of Jessie’s collection, and the skill and taste with which it is displayed. She’s featured some of these specimens on her Instagram, which I strongly recommend.

This is a Galeamopus, roughly two feet long, sculpted by James Herrmann (who also made the life-size Aquilops sculpture and bust) for the Cincinnati Museum Center.

Here’s what it looks like on the other side.

From behind.

And from the front.

I dig this. I’m sure someone else must have done this half-skeletal reconstruction, half-fleshed life restoration style of sculpture before, but I can’t think of any museum-quality examples. The bronze is a nice touch.

Here’s a convincingly chunky Allosaurus.

About the sculpting process, James wrote (in an email with permission to cite):

I worked on all of the museum pieces with Glenn Storrs, Ph.D., vertebrate paleontologist with the Cincinnati Museum Center. He would tell me what he envisioned and provide me with reference material, I would sculpt it, take the clay to Glenn for his critique, take it back and make revisions. We went through several cycles of this for each piece and when I received the final approval I took each piece to the foundry.

Tyrannosaurs are to museums what roller-coasters are to amusement parks. Here’s Daspletosaurus.

My favorite thing about these sculptures is why they’re done in bronze. It’s not just for posterity. James again:

The idea was to provide a small sculpture of each skeletal reconstruction on display for people to touch and feel. It was felt that this element of touch would be particularly important to accommodate the needs of the visually impaired museum visitor. I will feel like I have achieved success when the patina is rubbed off parts of the bronze.

One more, a life-size bust of Galeamopus.

In addition to having these on display at the Cincinnati Museum Center, James will be producing these sculptures as limited editions. If you’re interested, please visit http://www.herrmannstudio.com/.