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It’s World Penguin Day! Watch your back though… these penguins aren’t as nice as they seem. But they need us to be nice to them!

Hahaha?Whether you watch a classic GIF like the one above, or a kid-friendly TV/film documentary, you might get the impression that penguins lead carefree, or at least silly or slapstick, lives– happy feet and all that. It works for Hollywood: a Charlie Chaplin comedy relief role to play.  And that’s the vision of penguins I grew up with: they were living cartoons to me.

But what’s the reality? Plenty of documentaries, most notably to my mind the recent Attenborough’s “Frozen Earth” episodes or “March of the Penguins” film, have dealt with the darker side to these two-toned, tuxedo-toting antipodeans. And anyone who has experienced penguins in the wild has probably seen those not-so-light facets of penguinity firsthand. On realiizing just how compulsively horny young “hooligan cock” male penguins were, Natural History Museum ornithologist Douglas Russell wrote: ““just the frozen head of the penguin, with self-adhesive white O’s for eye rings, propped upright on wire with a large rock for a body, was sufficient stimulus for males to copulate and deposit sperm on the rock.”

Stomach-Churning Rating: 5/10; some tears may be shed over cute baby penguins and you might choke if you’re a rhea trying to swallow one, but the anatomy shown is mostly skeletal or dessicated. No penguin juices. Except those just mentioned above.

I’m quick to admit, I didn’t know much about penguins until recently. I couldn’t name many species or say much about their behaviour, anatomy or evolutionary history. When I was a graduate student at Berkeley, I was enthused by a now-classic, elegantly simple study (published in 2000) that fellow PhD student Tim Griffin and biomechanist Dr. Rodger Kram conducted on penguin waddling. They found that the waddling gait of penguins isn’t mechanically disadvantageous, as it appears, but rather is a way that they conserve energy while walking. It’s the short legs, instead, that make their gait metabolically expensive, because shorter legs mean that more frequent, costly steps need to be taken, incurring high costs due to rapid firing of leg muscles to support the body. My vicarious enjoyment of Griffin’s & Kram’s research began my scientific introduction to penguins. Fast forward to 2014: I get a crash course in penguinology.

Punta Tombo (4)

Mostly-fledged Magellanic penguin

That’s what this post is about, and how it brought me in touch with The Existentialist Penguin– the haggard, storm-tossed, predator-harried, starved and bullied wanderer of wastelands.

My personal introduction to penguins over the past year has been initiated by a collaboration with PhD student James Proffitt and long-time colleague Dr. Julia Clarke, both at the University of Texas in Austin. They kindly invited me to collaborate on applying modern biomechanics to the surprisingly excellent fossil record of penguins (Sphenisciformes), among other extant water birds. Before diving into it all, I happened to go to Argentina.

Punta Tombo (2)

Penguin tries to keep cool in the shade, opening its mouth to shed heat in the autumn sun.

Just before I travelled to Patagonia on unrelated business (to study sauropodomorph dinosaurs!), I did a little googling and came across Punta Tombo reserve, near the city of Trelew that I was visiting (more about that in a future post!). It’s where some 1+ million Magellanic penguins (Spheniscus magellanicus) gather every southern summer to breed and fledge before making a long ~5 month swim up to Brazil. I asked my host, Dr. Alejandro Otero, if we might take a day off to visit this spot, where guanacos, rheas and other wildlife were also said to be common, and he basically said “Hell yes!” as he’d never been there. My Flickr photostream gives a big set of my favourite photos from that trip, but here are some others below, to show some of my experiences. We rented a car and took a lovely 90-minute drive south across the Patagonian plains, observing wildlife like tinamous (yes! So exciting for me) as we went. You could get within 1.5m of the penguins according to park rules, and the penguins were very permissive of that!

This jaunty chap was staying put in his burrow while people walked by. We came closer and he kept rotating his head around, staring at us. I first took it as cute juvenile behaviour, but on later observations of penguins realized it was a threat- "My beak is sharp! Stay back, bro, or I'll glock ya!"

This jaunty chap was staying put in his burrow while people walked by. We came closer and he kept rotating his head around, staring at us. I first took it as cute juvenile behaviour, but on later observations of penguins realized it was a threat- “My beak is sharp! Stay back, bro, or I’ll glock ya!”

The video below shows a penguin encounter that left me with no doubts that these animals don’t mess around. The smaller penguin escaped, losing its cool burrow and some of its tough hide, too. Indeed, penguins can be remarkable assholes to each other.

With battles like this erupting all around us, where the penguins struggled to find shade in the desert-like inland parts of the park, often hundreds of meters away from the cool ocean, it came as no surprise to find casualties. The juveniles (and some remaining adults; most having left by now while the ~1 year-old juveniles fledge) not only battled, but also fasted, and roasted in the heat as they shed their insulatory fluff for waterproofed streamlining. This poor little flat Spheniscus had been trodden a bit past streamlined:Punta Tombo (3)

Near the end of our visit, just after I saw an informative sign about the lesser rhea or “choique” (Pterocnemia/Rhea pennata), we managed to get very close to a rhea and follow it for a while, as penguins stood around in apparent disinterest. I’ll never forget that meeting: two flightless birds, yet adapted to such different lifestyles and habitats. The penguins were in the rhea’s domain; a hot, wind-blown, scree-scoured scrubland on the edge of the fertile ocean.rhea-penguin

The choique soon found a dry old hatchling penguin carcass, no meatier than the surrounding thickets, and tried to swallow it. The loss of teeth by its distant ornithurine ancestors proved to be a bad move, because it struggled to get the jerky-like mass through its beak:

That Punta Tombo visit was an experience I’ll never forget. I returned to the UK, abuzz with excitement about penguins. I “got” them now, I felt, at least in a very unscientific, anthropomorphic way. It took the face-to-beak experience to drive that home, more than any emotive film treatment could. Whether enduring Antarctic wintery blasts or unforgivingly hot and dry, burrow-speckled coastal badlands, penguins are buggers with true grit. Survivors, as their >60 million year fossil record attests to. On my return, I delved through my photos of museum specimens to get a better appreciation for penguin anatomy, preparing to also get familiar with that fossil record; all as part of that ongoing work with Proffitt and Clarke. Here’s some of that anatomy:

My first encounter with a penguin in the wild is probably this specimen washed up on a beach in Uruguay. I'm going with the tentative ID of a juvenile penguin skeleton; probably Magellanic.

My first encounter with a penguin in the wild (but not a live one) is probably this specimen washed up on a beach in Uruguay. I’m going with the tentative ID of a juvenile penguin skeleton (short foot; flat wing bones); probably Magellanic. The bevy of vertebrate morphologists investigating dead penguins on this beach during our conference in 2010 will not soon be forgotten!

Magellanic penguin skeleton, "flying" through the Punta Tombo visitor centre.

Magellanic penguin skeleton, “flying” through the Punta Tombo visitor centre.

University Museum of Zoology Cambridge skeleton of one of the "great penguin" (do not confuse with the great pumpkin!) species; either King (patagonicus) or Emperor (forsteri).

University Museum of Zoology Cambridge skeleton of a “great penguin” (do not confuse with the great pumpkin!) species of Aptenodytes; either King (patagonicus) or Emperor (forsteri). Characteristic features, in addition to the robust, dense skeleton, include the short neck, flattened but robust wings and scapulae, robust furcula (wishbone), stubby legs (with a big blocky patella) and thin but longish tail (supposedly used to balance with while walking/standing).

I’ll visit some more penguin anatomy in coming images- those photos are just teasers. And they set the stage for me to go back to my one-stop-shopping for awesome ornithological specimens, the Natural History Museum at Tring (images below presented with kind permission from the Natural History Museum, London; but I took the photos), to pick up an assortment of 11 frozen penguins from helpful curator Hein van Grouw! Such as this “gagged” King penguin:
NHMUK penguin

And this handsome Emperor penguin, going through the Equine Imaging Centre’s CT scanner as I do my usual routine of (1) get cool critters, (2) barrage them with radiation to peek inside:penguin CT (3)

CT scanner monitors as I scan a penguin; mid-torso x-ray slice shown on the right.

CT scanner monitors as I scan a penguin; mid-torso x-ray slice shown on the right.

Awwwwww... baby Gentoo penguin (Pygoscelis papua). Unhappy feet, I'm afraid.

Awwwwww… baby Gentoo penguin (Pygoscelis papua— EDIT: Probably Aptenodytes; see comments below). Unhappy feet, I’m afraid… Happy CT scanning, however– specimens like this are NOT easy to come by in these northern nether regions!

Because I love the CT scan images of these penguins so much (their skeletons are awesome and bizarre!), I’ll share the pilot scans of the best ones now:

Calling all penguin experts! What's up with this? Is that really how much gastrolith volume a penguin carries, or did a museum curator stick rocks up its bum? Seems very caudal in position. I'm fascinated.

Calling all penguin experts! What’s up with this? Is that really how much gastrolith (stomach stone; near bottom of image) volume a penguin carries (answer after some literature reading: maybe yes!), or did a museum curator stick rocks up its bum? It seems very caudal in position, and this is consistent with other animals I’ve seen (some below). A paper on this phenomenon and potential role in ballast is here. Another here.

Side view.

Side view. Nice view of the head at least.

The fluffy baby shown in the photo above. Nice pose, and lots of anatomy shown. And check it out- gastroliths?!? In such a young animal-- is it even feeding yet?

Young juvenile. Nice pose, and lots of anatomy is shown. And check it out- gastroliths?!? In such a young animal– is it even feeding yet? (presumably straight after hatching) And they are relatively big pebbles, too! If I noticed this 5 years ago, it would have been a nice paper to report- first recognition of gastroliths in penguin chicks seems to have been then. Indeed, that study observed some chicks intentionally swallowing stones.

Another youngun.

Another youngun; the fluffy one from the photo above. More rocks up its wazoo.

Three wee little chicks.

Three wee little chicks, all with stomach stones.

CT reconstruction of adult skeleton. This specimen was gutted and flattened, so the gastroliths are few and scattered. Check out the long tail:

From recent skeletons to fossil ones, penguins have wacky anatomy; they break most of the “rules” of being a proper bird, putting other oddballs like rheas to shame. I can’t ably review the many penguin species we know of, but the ancient Palaeocene penguin Waimanu features prominently in recent scientific discussions of penguin evolution, such as the superb research and blog of Dan Ksepka  as well as many workers in the southern hemisphere. I haven’t had a chance to inspect that creature’s bones, but while in Trelew, Argentina, I was very pleased to run into some excellent specimens of a later animal:

Part of the rather nice skeleton of Palaeospheniscus patagonicus, an Oligocene/Miocene largish penguin; from the MFN collections in Trelew, Argentina and collected nearby.

Part of the nice skeleton of Palaeospheniscus patagonicus, an Oligocene/Miocene largish penguin; from the MEF collections in Trelew, Argentina and collected nearby. The genus has been known since Ameghino’s description in 1891, and is closely related to living penguins, especially Aptenodytes. It was not a large penguin, but at about 5kg body mass was no slouch as birds go (roughly similar in size to a Magellanic penguin). I also got to see  Madrynornis mirandus, a Miocene form.

For me, the diagnostic trait of a penguin skeleton: the very short, tobust tarsometatarsus. From Palaeospheniscus, as above.

For me, the diagnostic trait of a penguin skeleton: the very short, tobust tarsometatarsus. From Palaeospheniscus, as above. The great palaeontologist GG Simpson wrote of it: “Despite the innumerable variations in details, the tarsometatarsi, on which all species but P. robustus are based, are quite stereotyped in general structure and leave little doubt that the forms placed here by Ameghino do all belong to a natural group.” A ratio of length to proximal width of >2 is typical of most penguins.  Synapomorphy FTW!

From beach skeletons, to mass suffering of landbound birds, to 3D imaging and fossil skeletons, I’ve had quite the immersion in penguinness lately. And through that experience, I’ve been drawn closer to penguins in more ways than one. I’ve been impressed by their adaptability and durability. In some ways, penguins’ adaptations to harsh freezing winters in wastelands also aid them to survive harsh baking summers in dry badlands.

Yes, those badlands are still coastal, and penguins can still drink the saltwater and excrete salt via their supraorbital glands, but those penguins in Punta Tombo were not having a keg party. They were clearly enduring some serious discomfort, and not all making it through the ordeal. I watched silently along with other penguins as one penguin lay prone in an awkward pose on a bleached-white stretch of hardpan soil, while one flipper meekly raised, then flopped down. It was not long for this world, and there was a host of large scavengers around ready to make the most of that, while penguin-eating giant petrels (a sister group to penguins) wheeled overhead.

penguin-waddle

Waddlers of the wastes

While penguins still spend most of their lives at sea, they retain a sometimes astonishing array of behaviours they use on land: burrowing, hopping/jumping, costly short-legged (but efficiently waddling) walking, and perhaps more that we haven’t yet discovered! Their unique anatomy reflects a compromise between all these factors, and we’re fortunate to have knowledge of their fossil record that shows a lot of detail on how they evolved it all. While penguins are a highly aquatic species, they show how aquatic and terrestrial adaptations can coexist in harmony; it’s not just a black-or-white issue. But with climate change in progress, the ~18 species of penguins have some rapidly altering challenges to adapt to, or go the way of Waimanu. This is a critical Kierkegaardian moment for The Existentialist Penguin.

I raise a glass in toast to that versatile, resilient, gravel-gizzarded Existentialist Penguin! May it persevere all the troubles our ever-changing world throws at it, as it has done since the Palaeocene. And may we draw inspiration from its tenacity, to face our own troubles, together on this crazy spinning globe!

Cheers!

by animalloz, on deviantart

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And I post my blog and stare
Into x-rays of an ostrich
I’ve always known that radiographs never lie
People always say “that’s cool”
To see x-rays of an ostrich
So keen to know what
Lies behind the skin

(evolved from “Eyes of A Stranger” by Queensrÿche, from the epic masterpiece of Operation: Mindcrime (1988). One of my favourite albums of all time, and a fantastic concept album too. The band was operating at their peak. Tight! Drug addict Nikki gets brainwashed by the evil Dr. X and made to assassinate a nun, Sister Mary, who was a prostitute, and then there’s like a revolution or something, and things get all screwed up and no one ends up happy – or alive. All the while, Geoff Tate is singing his guts out. Anyway, I got to see them play the whole album live in 1990 in Madison, WI, for the filming of Operation: Livecrime, which was like a Mecca moment for me back then. Look for me (pre-bald years) in about the 6th row. )

What does that album have to do with the number 2 (two days left in Freezermas)? Hmm… Track 2 is the instrumental Anarchy-X, and today’s post is about X-rays as well as that funky ostrich (2 legs good! 2 toes good, too!) again, so I’m satisfied, and by this point you’re probably just oggling the mind-blowing images below anyway, so fuck it!

Stomach-Churning Rating: 2/10; just X-rays.

Tech/MRes Kyle Chadwick, Renate Weller and the equine imaging team at the RVC took these x-rays of our birdie for us and for an artist who is doing a big x-ray animal art show (more news on this soon!)– thanks to all of them for some truly awesome images! I could stare at the intricate details in these images for hours– go ahead, do it. Click to emostrichinate them (this post needs to be viewed on nice big screen), and oggle away…

Head and neck.

Head and neck.

Another view of the same.

Another view of the same. The highly flexible esophagus and trachea can be seen going diagonally across the neck; twisting from ventral to dorsal. It’s floppy, so it can do that.

Neck near the head; tapering.

Neck near the head; tapering.

Middle of neck. Check out the rings of the trachea!

Middle of neck. Check out the rings of the trachea!

Base of neck and shoulder

Base of neck and shoulder.

Shoulder and chest. Hard to image; thick and dense (still was frozen).

Shoulder and chest. Hard to image; thick and dense (still was frozen), hence the whiteout toward the left side of the image.

Check out that wing!!

Check out that wing!!

Ankle- note the big calloused pad that ostriches rest on (right side of image).

Ankle- note the big calloused pad that ostriches rest on (right side of image).

That two-toed foot... but did you know that normally the missing 2nd toe is still there as a fibrous remnant on the 3rd toe?

That two-toed foot… but did you know that normally the missing 2nd toe is still there as a fibrous remnant on the 3rd toe?

Tomorrow: the final day of Freezermas. What will it be?

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Freezermas continues! Today we have a treat for you. Lots of detailed anatomy! This post comes from my team’s dissections of an ostrich last week (~3-7 February 2014), which I’ve been tweeting about as part of a larger project called the Open Ostrich.

However, before I go further, it’s as important as ever to note this:

Stomach-Churning Rating: 9/10: bloody pictures of a dissection of a large ostrich follow. Head to toes, it gets messy. Just be glad it wasn’t rotten; I was glad. Not Safe For Lunch!

If the introductory picture below gets the butterflies a-fluttering in your tummy, turn back now! It gets messier. There are tamer pics in my earlier Naked Ostriches post (still, a rating of 6/10 or so for stomach-churning-ness there).

All photo credits  (used with permission) on this post go to palaeoartist Bob Nicholls (please check out his website!), who got to attend and get hands-on experience in extant dinosaur anatomy with my team and Writtle College lecturer Nieky VanVeggel (more from Nieky soon)!

Research Fellow Jeff Rankin, myself and technician/MRes student Kyle Chadwick get to work.

Research Fellow Jeff Rankin, myself and technician/MRes student Kyle Chadwick get to work, removing a wing.

This is a male ostrich, 71.3 kg in body mass, that had gone lame in one foot last summer and, for welfare reasons, we had to put down for a local farmer, then we got the body to study. We took advantage of a bad situation; the animal was better off being humanely put down.

The number for today is 6; six posts left in Freezermas. But I had no idea I’d have a hard time finding a song involving 6, from a concept album. Yet 6 three times over is Slayer’s numerus operandi, and so… The concept album for today is Slayer’s  1986 thematic opus “Reign in Blood” (a pivotal album for speed/death metal). The most appropriate track here is the plodding, pounding, brooding, then savagely furious “Postmortem“, which leads (literally and figuratively, in thunderous fashion) to the madness of the title track, after Tom Araya barks the final verse:

“The waves of blood are rushing near, pounding at the walls of lies

Turning off my sanity, reaching back into my mind

Non-rising body from the grave showing new reality

What I am, what I want, I’m only after death”

I’m not going to try to reword those morbid lyrics into something humorous and fitting the ostrich theme of this post. I’ll stick with a serious tone for now. I like to take these opportunities to provoke thought about the duality of a situation like this. It’s grim stuff; dark and bloody and saturated with our own inner fears of mortality and our disgust at what normally is politely concealed behind the integumentary system’s viscoelastic walls of keratin and collagen.

But it’s also profoundly beautiful stuff– anatomy, even in a gory state like this, has a mesmerizing impact: how intricately the varied parts fit together with each other and with their roles in their environment, or even the richness of hues and multifarous patterns that pervade the dissected form, or the surprising variations within an individual that tell you stories about its life, health or growth. Every dissection is a new journey for an anatomist.

OK I’ve given you enough time to gird yourself; into the Open Ostrich we go! The remainder is a photo-blog exploration of ostrich gross anatomy, from our detailed postmortem.

(more…)

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Here, I give you a long-planned post on the patella (“kneecap bone”) of birds, which was my Royal Society Senior Research Fellowship sabbatical project for 2012-13. This is only a brief introduction to the anatomical issues at hand, err, I mean at knee…

Stomach-Churning Rating: 6/10; mostly skeletons/fossils, but there are a few images of the dissection of a guineafowl, which is fresh and meaty.

Archaeopteryx, the Berlin specimen. Helluva fossil, but nary a patella!

Archaeopteryx, the Berlin specimen. Helluva fossil, but nary a patella!

The question I am exploring, first of all, is simply how the patella evolved, because it seems to be present in almost all living birds. However, it is absent in all non-avian dinosaurs, and indeed most Mesozoic birds, too. There is barely a hint of any precursor structure (a “patelloid”) in other reptiles, but lizards evolved their own patella that is quite different (a flattened lozenge, not a rectangular structure lying tightly confined in a “patellar groove” on the femur as it is in birds). Mammals evolved the knobbly, hemispherical kind of kneecap that we’re familiar with, possibly on several occasions (a different story!). So the patella evolved at least three times in the lizard, mammal and bird lineages– and possibly more than once in each of these groups. And that’s about it for almost 400 million years of tetrapod evolution, except for a few very rare instances in fossils and sort-of-patella-like things in some frogs or other weirdos.

Fossil birds exhibit no clear presence of a patella until we come very close to modern birds on the avian stem of the tree of life (see below). And then, suddenly in modern birds, there is a lot of variation and not much good documentation of what kind of patella exists. This makes it challenging to figure out if the patella is ancient for modern birds or if it evolved multiple times, or how it changed after it first evolved– let alone bigger questions of what the patella was “for” (performance benefits, functional consequences, etc.; and developmental constraints) in the birds that first evolved it.

Considering that the patella is such an obvious bone in some birds, and certainly affects the mechanics of the knee joint (forming a lever for the muscles that cross it; homologous to our quadriceps muscles) and hence locomotion, it is a compelling research topic for me.

What follows is a pictorial guide to the patella of some birds, in sort of an evolutionary/temporal sequence (see my earlier post for a recap of some major groups), with a focus on animals I’ve studied more intensively so far (with >10,000 species, there is a lot that could be done):

Gansus, IVPP V15080
The early Cretaceous bird Gansus (from the IVPP in Beijing), represented by many beautifully preserved specimens, all of which lack a patella. This absence is characteristic of other stunningly preserved fossil Chinese birds, indicating that this is almost certainly an ancestral absence of a patella, until…

The famed Cretaceous diving (flightless) bird Hesperornis, from Wikipedia/Smithsonian.  Note the massive, conical/crested patella in front of the  knee (jutting up and overlapping the ribs/vertebrae close to the pelvis; see also below). That elongate patella is characteristic of many diving birds that use foot-propelled swimming; it has evolved many times in this fashion. Other hesperornithiform birds show some transformational states in their anatomy toward this extreme one.
Hesp-patella

Check this out! More Hesperornis (cast), with the femur on the left and the patella on the right. The bloody patella is almost as long as the femur! That’s nuts. With kind permission from the Natural History Museum, London.

Exhibited ostrich skeleton in left side view showing the patella (white arrow).
Exhibited ostrich (Struthio camelus) skeleton in left side view showing the patella (white arrow), on exhibit atThe Natural History Museum at Tring, Hertfordshire, UK. Ostriches are remarkable in that they have this elongate patella (actually a double patella; there is a smaller, often-overlooked second piece of bone) and yet are rather basal (closer to the root of the modern avian family tree)– however, they obviously are specialized in ways other than this double patella, most notably their very large size, flightlessness, and elongate legs. So the unusual patella is more likely linked to their odd lifestyle than a truly primitive trait, at least to some degree (but stay tuned: what happened with the patella in other members of their lineage, the ratites/palaeognaths, is much less well understood!).
Note that ostriches and Hesperornis together hint that the presence of a patella might have been an ancestral trait for living birds, but their patellae are so different that the ancestral state from which they evolved must have been different, too; perhaps simpler and smaller. Hence we need to look at other birds…
Skinned right leg of guineafowl, Numida meleagris.
Skinned right leg of a Helmeted Guineafowl, Numida meleagris, above. That whitish band of tissue in the middle of the screen, on the front of  the knee, is part of what is concealing the patella. That is an aponeurosis (connective tissue sheet, like a thin tendon) of the muscles corresponding to our “quads” or our tensor fascia latae, detailed more below. Guineafowl are fairly basal and well-studied in terms of their bipedal locomotion, so they are an important reference point for avian form and function.
Right guinefowl leg, with patella semi-exposed.
Right guineafowl leg, with patella exposed. Here I’ve peeled away that white band of tissue  and associated muscles, which have been reflected toward the bottom of the screen (AIL and PIL labels corresponding to the anterior and posterior parts of the Iliotibialis lateralis muscle). The tip of the scalpel is contacting the patella. It’s not much to see, but lies atop the bright yellow fat pad that cushions it against the femur. You should be able to see a groove in the end of the femur just above that fat pad, which is where the patella sits and slides up and down as the knee moves/muscles contract. This is called the patellar groove, or sulcus patellaris.
Left leg of a guineafowl (with right tibiotarsus behind it) showing both patellae in articulation; in medial (inside) view. The  patella is the little rectangular bit of bone in the top middle of the screen, interposed between femur and tibiotarsus.
Left leg of another guineafowl (with right tibiotarsus behind it, on the left) showing the patellae in articulation; in medial (inside) and cranial (front) views, respectively. The patella is the little rectangular bit of bone in the top middle of the screen, interposed between femur (thigh) and tibiotarsus (shank). With kind permission from the Natural History Museum, London.  
Penguin-patella
Right leg of a Cape Penguin (Spheniscus demersus) from the University Museum of Zoology in Cambridge, showing the big lumpy patella in this wing-propelled diver. They still walk long distances on land, so presumably a patella plays some role in their gait, helping to explain its large size, which like the ostrich and Hesperornis seems to be a novel trait. Notice the groove across the patella, made by the tendon of the ambiens (like our sartorius/”tailor’s muscle”), which crosses from the inside to the outside of the leg via this route. This groove is often considered a useful phylogenetic character in modern birds, as its contact with the patella (sometimes via a hole, or foramen) varies a lot among species.
Buceros skeleton UMZC
A hornbill, Buceros sp., from the UMZ Cambridge museum as well. This displays the possibly-more-typical, little rounded patellar nubbin that many birds have. See below for more.

Buceros knee closeupCloseup of the knee/patella of the hornbill, Buceros sp., from above. Not much to squawk about, patella-wise, but it’s there.

And so we complete our quick tour of the avian patella, in its grand variation and humble beginnings.

Why does an ostrich have a patella and a Tyrannosaurus, Edmontosaurus or Triceratops did not? Why were birds the only bipedal lineage to evolve a patella (mammals and lizards gained a patella as small quadrupeds), and why did some bipeds like kangaroos “lose” (reduce to fibrous tissue, apparently) their patella?

These are the kinds of mysteries my group will now be tackling, thanks to a generous Leverhulme Trust grant on sesamoid bone ontogeny, mechanics and evolution.  My group is now Dr. Vivian Allen as the postdoc, Sophie Regnault as the PhD student, and Kyle Chadwick as the technician and MRes student, along with numerous collaborators and spin-off projects. We’re looking forward to sharing more! But for now, I hope that I’ve engendered some appreciation for the avian patella, as the silly title indicates (“fella” used in the general sense of anyone!). This work is all unpublished, but some of this should be out in not too long, in much more lavish detail! Much as the patella is the “forgotten lever “of the avian hindlimb, it is the fulcrum about which a substantial part of my research group’s activity now pivots.

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A photo blog post for ya here! I went to Dublin on a ~28 hour tour, for a PhD viva (now-Dr Xia Wang; bird feather/flight evolution thesis) earlier this month. And I made a beeline for the local natural history museum (National Museum of Ireland, Natural History building) when I had free time. So here are the results!

Stomach-Churning Rating: Tame; about a 1/10 for most, but I am going to break my rule about showing human bodies near the end. Just a warning. The bog bodies were too awesome not to share. So that might be 4/10-8/10 depending on your proclivities. They are dry and not juicy or bloody, and don’t look as human as you’d expect.

Simple Natural History museum entrance area.

Simple Natural History museum entrance area.

Adorable frolicking topiaries outside the NHM.

Adorable frolicking topiaries outside the NHM.

Inside, it was a classical Victorian-style, dark wood-panelled museum stuffed with stuffed specimens. It could use major refurbishment, but I do love old-fashioned exhibits. Get on with it and show us the animals; minimize interpretive signage and NO FUCKING INTERACTIVE COMPUTER PANELS! So by those criteria, I liked it. Some shots of the halls: hall2 hall1 hall3 hall4 hall5 hall6 And on to the specimens!

Giant European deer ("Irish elk"). I looked at these and thought, "why don't we see female deer without antlers ever? then noticed one standing next to these; photo was crappy though. :(

Giant European deer (“Irish elk”). I looked at these and thought, “why don’t we see female deer without antlers ever? then noticed one standing next to these (you can barely see it in back); too bad my photo is crappy.

Superb mounted skeleton of giraffe (stuffed skin was standing near it).

Superb mounted skeleton of giraffe (stuffed skin was standing near it).

A sheep or a goat-y thingy; I dunno but it shows off a nice example of the nuchal ligament (supports the head/neck).

A sheep-y or a goat-y beastie; I dunno but it shows off a nice example of the nuchal ligament (supports the head/neck).

Yarr, narwhals be internet gold!

Yarr, narwhals be internet gold!

Giant blown glass models of lice!

Giant blown glass models of lice!

Who doesn't like a good giant foramanifera image/models? Not me.

Who doesn’t like a good giant foramanifera image/model?

"That's one bigass skate," I murmured to myself.

“That’s one bigass skate,” I murmured to myself.

"That's one bigass halibut," I quipped.

“That’s one bigass halibut,” I quipped.

Tatty basking shark in entry hall.

Tatty basking shark in entry hall.

Irish wolfhound, with a glass sculpture of its spine hanging near it, for some reason.

Irish wolfhound, with a glass sculpture of its spine hanging near it, for some reason.

Stand back folks! The beaver has a club!

Stand back everyone! That beaver has a club!

Skull of a pilot whale/dolphin.

Skull of a pilot whale/dolphin.

Nice anteater skeleton and skin.

Nice anteater skeleton and skin.

Nice anteater skeleton and skin.

Nice wombat skeleton and skin.

Sad display of a stuffed rhino with the horn removed, and signage explaining the problem of thefts of those horns from museum specimens of rhinos worldwide.

Sad display of a stuffed rhino with the horn removed, and signage explaining the problem of thefts of those horns from museum specimens of rhinos worldwide.

But then the stuffed animals started to get to me. Or maybe it was the hangover. Anyway, I saw this…
creepy proboscis (1) creepy proboscis (2)

A proboscis monkey mother who seemed to be saying “Hey kid, you want this yummy fruit? Tough shit. I’m going to hold it over here, out of reach.” with a disturbing grimace. That got me thinking about facial expressions in stuffed museum specimens of mammals more, and I couldn’t help but anthropomorphize as I toured the rest of the collection, journeying deeper into surreality as I progressed. What follows could thus be employed as a study of the Tim-Burton-eseque grimaces of stuffed sloths. Click to emslothen.

sloths (1) sloths (5)sloths (4) sloths (3) sloths (2)

Tree anteater has a go at the awkward expression game.

Tree anteater has a go at the awkward expression game.


This completed my tour of the museum; there were 2 more floors of specimens but they were closed for, sigh, say it with me… health and safety reasons. Balconies from which toddlers or pensioners or drunken undergrads could accidentally catapult themselves to their messy demise upon the throngs of zoological specimens below. But the National Museum’s Archaeology collection was just around the block, so off I went, following whispered tales of bog bodies. There will be a nice, calm, pretty photo, then the bodies, so if peaty ~300 BCE cadavers are not your cup of boggy tea, you can depart this tour now and lose no respect.

Impressive entrance to the National Museum's Archaeology building.

Impressive entrance to the National Museum’s Archaeology building.

The bog bodies exhibit is called “Kingship and Sacrifice“. It is packed with cylindrical chambers that conceal, and present in a tomb-like enclosed setting, the partial bodies of people that were killed and then tossed in peat bogs as honoraria for the ascension of a new king. The peaty chemistry has preserved them for ~2300 years, but in a dessicated, contorted state. The preservation has imparted a mottled colouration and wrinkled texture not far off from a Twix chocolate bar’s. Researchers have studied the bejesus out of these bodies (including 3D medical imaging techniques) and found remarkable details including not just wounds and likely causes of death (axes, strangling, slit throats etc) but also clothing, diet, health and more.

Here they are; click to (wait for it)… emboggen:

BogBodies (1) BogBodies (2) BogBodies (3) BogBodies (4) BogBodies (5) BogBodies (6)

Did you find the Celtic armband on one of them?

Finally (actually this happened first; my post is going back in time), I visited UCD’s zoology building for the PhD viva and saw a few cool specimens there, as follows:

Giant deer in UCD zoology building foyer.

Giant deer in UCD zoology building foyer, with a lovely Pleistocene landscape painted on the wall behind it.

Sika deer in awkward posture in Univ Coll Dublin zoology building's foyer.

Sika deer in an awkward posture (what is it supposed to be doing?) in Univ Coll Dublin zoology building’s foyer.

The pose of this ?baboon? struck me as very peculiar, and menacing- reminiscent of a vampire bat's pose, to me.

The pose of this ?baboon?mandrill struck me as very peculiar and menacing- reminiscent of a vampire bat’s pose.

A whole lotta chicken skeletons in a UCD teaching lab.

A whole lotta chicken skeletons in a UCD teaching lab.

After the viva we went out for some nice Chinese food and passed some Dublin landmarks like this:

Trinity College entrance, I think.

Trinity College entrance, I think.Former Irish Parliament; now the Bank of Ireland.

And we wandered into a very posh Irish pub called the Bank (on College Green), which displayed this interesting specimen, as well as some other features shown below:

Replica of illuminated old Gaelic manuscript.

Replica of illuminated 9th Century gospel manuscript “The Book of Kells”, with gorgeous Celtic art.

Vaults near toilets in the Bank pub.

Vaults near toilets in the Bank pub. Almost as cool as having giant freezers down there.

Nice glass ceiling of the Bank pub.

Nice glass ceiling of the Bank pub.

And Irish pub means one big, delicious thing to me, which I will finish with here– much as I finished that night off:

Ahhh...

Ahhh… ice cold.

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Less words, more pictures in this post, and I’ll get the one lame cake joke out of the way early. I’ve nearly finished my research blitz through the postcranial material of the NHM-Tring’s osteological collection and have made some pit-stops for cake skulls now and then when I see one that pleases me. Now I shall present a survey of some of the species I’ve examined. I’ll proceed up from the base of the crown clade of living birds (Neornithes/Aves; the most recent common ancestor of living birds and all its descendants) and first take a tour of Palaeognathae; the ratites and kin; then move another step up into the Neognathae, first featuring the lineage featuring the ground fowl (Galliformes) and then the waterfowl (Anseriformes). If all this taxonomy and phylogeny is a bit much, check out this page for a brush-up on the bushy branches of bird biodiversity.

First, lots of bones of our cast of currasows, chachalacas, cassowaries and other kooky characters. And then, perhaps, a stop to the excessive alliteration. Finally, I will finish with some examples of species oddity (hat tip to Chris Hadfield).

Stomach-Churning Rating: 2/10- some bony pathologies but still just dry bones. Minimal cake jokes, and no filthy swearing this time.


BRING ON THE BONES:

My photographs are shown with kind permission from the Natural History Museum, London.

Exploded skull of an ostrich/ This takes skill.

Exploded skull of an ostrich, Struthio camelus. This kind of careful preparation takes crazy skill, and creates a thing of rare beauty.

Neat skull of a cassowary, Casuarius casuarius.

Imposing skull of a cassowary, Casuarius casuarius, with a rather worn head casque.

Mummified Owen's Little Spotted kiwi, Apteryx owenii.

Mummified Owen’s Little Spotted Kiwi, Apteryx owenii. The feathers were still soft and fluffy, but I would not call this specimen cuddly.

Dorsal view of the back/hips of the Great Spotted kiwi, Apteryx haasti.

Dorsal view of the back/hips of the Great Spotted Kiwi, Apteryx haasti. I like this photo and am not sure why. The symmetry and shading pleases me, I guess.

Front view of the back/hips of the Great Spotted kiwi, Apteryx haasti.

Front view of the back/hips of the Great Spotted Kiwi, Apteryx haasti, watching over my laptop and watching me while I write this blog on my laptop… so meta(ornithine)!

Wing of a kiwi, showing the fragile bones and feather attachments.

Wing of a kiwi, showing the fragile bones and feather attachments. “Apteryx” = “no wings”… well not quite. Click to emkiwi(?) so you can identify the individual bones, from the humerus right down to the fingers! I love this specimen.

The left leg (in front view) of the elephant-bird, Aepyornis maximus, from Madagascar, with a small moa nearby in left side view.

The titanic left leg (in front view) of the Elephant Bird, Aepyornis maximus, from Madagascar, with a small moa nearby in left side view. There’s so much awesomeness about elephant birds I don’t know where to start, but this is one good place to do so.

Mummified Unulated tinamou, Crypturellus undulatus.

The smaller end of the palaeognath scale: a mummified Undulated Tinamou, Crypturellus undulatus. Somehow the head got stuck into the abdominal cavity underneath the sternum, so this tinamou almost had its head up its arse. A tinamou with head in its proper position looks and sounds like this (video).

And now we take a left turn into the Galloanseres, most basal branch of the neognath birds, to see some of the neglected, strange early branches off from the “main line” that led to the modern diversity of ducks, geeses and swans (Anatinae, Anserinae).

Screamers (Anhimidae) are to Anseriformes as megapodes (see below; brush turkeys) are to Galliformes. By that I mean that both screamers and megapodes are very early branches off the main line of their respective lineages’ evolution, and both are quite strange when seen in that context… an unfair one, frankly; over-focused on the most familiar, “modern” or most speciose group. More about this issue further below.

This was my first hands-on experience with screamer anatomy; I was familiar from reading Tetrapod Zoology and other material about them. Check out the sound that gives them their name here! I’m now a big fan- they have so many strange features: oddly chunky but often very light bones, big feet with long toes, and then these switchblade-wrists, which would make Batman jealous:

Crested screamer, Chauna torquata, showing the wicked spur on the carpometacarpus.

Crested Screamer, Chauna torquata, showing the wicked spur (and smaller one) on the carpometacarpus.

Horned screamer, Anhima cornuta; similar carpometacarpal spur as in Chauna.

Horned Screamer, Anhima cornuta; similar carpometacarpal spurs as in Chauna.

Torso of a screamer seen in top view. Nice narrow body.

Torso of a screamer seen in top view. Nice narrow body, and no uncinate processes (spur-like bony struts that cross the ribs and act as levers for the muscles that move the ribcage during breathing)

The long, gracile, clawed toes of a screamer.

The long, gracile, clawed toes of a screamer. Those toes, especially as they belong to an animal called a screamer, are spooky for me. Note also: very little toe-webbing for a “waterfowl.”

Not to be outdone, on the Galliformes side of Galloanserae, we have some funky headgear in the Maleo (a megapode bird/Megapodiidae; a very basal branch of “brush turkeys” and kin) and curassows (part of the Cracidae; odd South American birds whose males make booming sounds, presumably using their head-casques as resonating chambers?):

Skull of a male maleo, Macrocephalon maleo.

Skull of a male Maleo, Macrocephalon maleo. AR Wallace famously pursued it, and here is its funky call.

Australian brush-turkeys, Alectura lathami i, at the Alma Park Zoo near Brisbane, Australia; they run wild there. Here they are doing what they are best known for: making a mound-like nest.

Australian brush-turkeys, Alectura lathami, at the Alma Park Zoo near Brisbane, Australia; they run wild there. Here they are doing what they are best known for: making a mound-like nest. We were doing kangaroo biomechanics experiments and they were everywhere. I was in awe to see such exotic (to me) birds; locals seemed not so enthused (the birds are loud and make a lot of mess).

Skull of Helmeted curassow, Crax/Pauxi pauxi.

Skull of Helmeted Curassow, Crax/Pauxi pauxi,  showing that resonating chamber. Along with this boom-boom-room, the male uses a piece of food that he holds to draw in the female; if she takes it, then it’s sexy time.

Foot of a Russian Black Grouse, Tetrao tetrix (nothing to do with a certain videogame), with and without flesh.

Foot of a Siberian Black Grouse, Tetrao tetrix (nothing to do with a certain videogame), with and without flesh. Regard the broad, feathered feet, well insulated and with plenty of surface area for prancing around in the snow or moorlands. Tetrao engage in a cool display pattern called lekking, in which the males group together and show off to watching females.

A theme in the section above that is not to be missed is that there is some amazing disparity of anatomical forms in these basal lineages of poultry-relatives. Don’t dismiss the Galloanserae as just boring food-birds! Heaps of not-so-well-studied species exist here, surely with a treasure trove of cool neontological and evolutionary questions waiting for the right person to ask! Darwin’s chickens may get their share of neglect, but that pales in comparison to how little we understand about many basal Galloanserae.

What a lot of people think of as a “ground fowl” or galliform way of life is more of a way of life somewhat typical of the Phasanidae- chickens, pheasants and their familiar kin. Megapodes, curassows, guans, grouse and other Galliformes do not necessarily do things in the “typical” ground fowl way, much as the earlier branches of the Anseriformes don’t always look/act like “proper water fowl” (i.e. Anatidae). The phenomenon at play here is one of the great bugaboos in biology: essentialism– the often implicit misconception that variation away from some abstract ideal is negligible, uninteresting or just not conceivable due to mental blinders. When we say something like “the chicken is a fascinating species” we are sliding down the essentialistic slope. There is no “the chicken.” Not really. Oh dear, speaking of slippery slopes, I’d best stop here before I start talking about species concepts. And no one wants that to happen! Anyway, essentialism still pervades a lot of modern scientific thinking, and has its place as a conceptual crutch sometimes. But in biology, essentialism can be very insidious and misleading. It burrows in deep into the scientific mind and can be hard to root out. Unfortunately, it is entrenched in a lot of science education, as it makes things easier to teach if you sweep aside the exceptions to the essentialist “rules” in biology. I catch myself thinking in static, essentialist ways sometimes. The punishment is no cake for a week; so awful. :)

And speaking of “normal” or “typical,” morphology is of course often not that way even within a species, age class or gender. Pathology is a great example; by definition it is abnormal. It is a shattering of the “essence” of animals, brought on by some malady.

Next I’ve highlighted some of the amazing pathologies I’ve seen in the Tring skeletons. There have been so many I’ve been unable to keep track of them– some of these birds had the stuffing beaten out of them, and I’m not talking about Thanksgiving turkeys. Some were captive animals, in which the pathology might be blamed on living an inappropriate environment, but some were wild-caught — given the extreme pathologies, it’s a wonder those even survived to be found, but perhaps less a surprise that they were caught.


BONES GONE BONKERS:

View of left knee of a specimen of the Highland guan, Penelopina nigra, showing some nasty osteoarthritis around the whole joint.

View of left knee of a specimen of the Highland Guan, Penelopina nigra, showing some nasty osteoarthritis around the whole joint. Eew.  A happier Guan sounds like this.

Femora and tibiae of the Blue-throated Piping Guan, Aburria cumanensis. Amazing pathology involving the left femur (broken, rehealed) and tibiotarsus (secondary infection?).

Femora and tibiotarsi of the Blue-throated Piping Guan, Aburria cumanensis. Amazing pathology involving the left femur (broken, rehealed) and tibiotarsus (secondary infection?). Interestingly, the non-fractured limb also showed some pathology, perhaps indicating general infection and/or arthritis in reaction to the severe damage to the other leg, or just increased load-bearing on that leg.

Little Chachalaca, Ortalis motmot, showing a broken and rehealed right femur and the tibiotarsus.

Little Chachalaca, Ortalis motmot, showing a broken and rehealed right femur and the tibiotarsus. As in the guan above, this animal was not walking for many weeks; its femur had snapped in two, but somehow melted back together. The tibiotarsus didn’t look too great, either; lumpy and bendy. In better times, the Chachalaca does the cha-cha like this.

These two specimens blew my mind. On the right is a normal Tetrao tetrix (Black grouse); on the left is one hybridized with another (unknown) species.

These two specimens blew my mind. On the left is a normal Tetrao tetrix (Black Grouse); on the right is one hybridized with another (unknown) species.

In the picture above, what amazed me first was the very unusual flattened pelvis/synsacrum of Tetrao, which characteristically is light and wide. But in the hybrid this morphology was completely gone; the pelvis had a more standard “galliform” (read: Phasianid)-like shape, deeper and narrower and more solid in build. I am guessing that the hybrid was a cross with a pheasant like Phasianus itself, whose anatomy would be more like this. Somewhere in here there is a fantastic evo-devo/morphometrics project waiting to happen.

That’s my quick specimen-based tour of “basal birds”. Beyond these two clades of Palaeognathae and Galloanseres, there lies the forebidding territory of Neoaves: much of living avian diversity, and extremely contentious in its phylogenetic relationships. I’m tackling them next for my research on the evolution of the patella/kneecap. But first, I’ll be at the NHM-Tring today for a whirlwind tour through the respectably speciose “normal” Galloanseres clades of Phasianidae and Anserinae+Anatidae, so off I go! (It’s my wife’s birthday celebration, so cake may have to wait for later this time)

So what do you think? What’s your favourite neglected “primitive” bird group (more apropos: early branching avian lineage that may still be very specialized, rare and poorly understood), or cool factoid about palaeognaths and basal neognaths?

No quaggas were harmed during the writing of this post.

No quaggas were harmed during the writing of this post. Polly wanna quagga?

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Bovids to the right of me, pinnipeds above, what's a guy to do but squee?

Bovids to the right of me, pinnipeds above, what’s a guy to do but squee?

I’ve been doing some osteological studies of the patella (bone in the major tendon in front of the knee; termed a sesamoid) that have included frequent visits to the Natural History Museum’s avian skeleton collection at Tring. It’s a cute little town, northeast of London, in the green county of Hertfordshire where I live and work. The museum at NHM-Tring is a great old school multi-storey display packed with skeletons and stuffed animals in dark wood cabinets, with many critters hanging from wrought iron railings or other suspensions above (see above). I blogged about the Unfeathered Bird exhibit (and book) that just finished up its tour there yesterday. And I’ll be blogging later, as I keep promising, about the cool things I’ve learned during the past year of my studies of the form, function, development and evolution of the patella.

As an aside, I heartily recommend doing research at the NHM-Tring. It’s away from the bustle (and arduous Tube trip) of the South Kensington NHM, and the curatorial staff are immensely helpful… and there is something else that makes the trip even more enjoyable, but you must read more below to find out about it.

Stomach-Churning Rating: 2/10; 150-year-old dry bones. But an advance warning to (1) diabetics and (2) pun-haters, for reasons that will become evident.

Dr Heather Paxton and Dr Jeffrey Rankin, postdoc researchers working on our collaborative BBSRC chicken biomechanics grant (see thechickenofthefuture.com), explain their science to an attentive Darwin.

Dr Heather Paxton and Dr Jeffery Rankin, postdoc researchers working on our collaborative BBSRC chicken biomechanics grant (see thechickenofthefuture.com), use the Structure & Motion Lab whiteboard to explain their science to an attentive Darwin.

Today I have a short pictorial exhibit of something wonderful I ran into while patellavating in the NHM collections. As often happens while doing museum research, I had a serendipitous encounter with a bit of history that blew my mind a little, and had me geeking out. These things happen because museum collections are stuffed with specimens that, to the right eyes or the right mindset, pack a profound historical whallop. As a scientist who is pretty keen on chickens (Gallus gallus), there are probably no museum specimens of chickens that would get me more excited about than the chickens Darwin studied in his investigations of artificial selection. In fact, most museum specimens of domestic chickens would not be that interesting to me, especially after seeing these ones.

Darwin wielded the analogy between artificial selection and his conceptual mechanism of natural selection in the first ~4 chapters of On the Origin of Species to clobber the reader with facts and try to leave them with no doubt that, over millennia, nature could craft organisms in vastly more complex and profound ways than human breeders could mould them over centuries. While people most often speak of Darwin’s pigeons when referring to Darwin and avians or artificial selection and variation, his chickens appear in The Origin and other writings quite often, too (most prominently, The Variation of Animals and Plants Under Domestication in 1868– more about that here). For example, from my 1st edition facsimile of The Origin from Harvard University Press, pp. 215-216:

Natural instincts are lost under domestication… It is not that chickens have lost all fear, but fear only of dogs and cats, for if the hen gives the danger-chuckle, they will run… and conceal themselves in the surrounding grass or thickets; and this is evidently done for the instinctive purpose of allowing, as we see in wild ground-birds, their mother to fly away. But this instinct retained by our chickens has become almost useless under domestication, for the mother-hen has almost lost by disuse the power of flight.”

Well told, Mr D!

I am also reminded of how chickens and Darwin have had darker relationships, such as this sad story. Or how evolution via Darwinian mechanisms crosses paths with pop culture in fowl ways, such as how tastes-like-chicken evolved, or how some say that chickens, over great periods of time, have been naturally selected in such a way that they are now heritably predisposed to cross roads, or that the amniote egg preceded the evolution of the genus Gallus by some 325+ million years. I see I am drifting and drifting further away from the topic at hand, so let me segue back to Darwin’s chickens. We’ll take this corridor there:

Inside the avian osteology collection at Tring. Sterlie at it might seem, places like this are  fertile breeding grounds for scientific discovery.

Inside the avian osteology collection at Tring. Sterile at it might outwardly seem, places like this are fertile breeding grounds for scientific discovery. And a sterile-looking collection means well cared-for specimens that will persevere for future discoveries.

So anyway, when museum curator Jo Cooper said to me something like “I have some of Darwin’s chickens out over on the other counter, do you want to have a look or shall I put them away?” my answer was quick and emphatic. YES! But only after lunch. I was hungry, and nothing stops me from sating that hunger especially when the sun is out and there are some fine pubs within walking distance! I settled on the King’s Arms freehouse, and had a delicious cheeseburger followed by a spectacularly good apple-treacle-cake with ice cream, expediently ingested while out on their sunny patio. Yum! I cannot wait to have that cake again. What a cake! Darwin’s bushy eyebrows would have been mightily elevated by the highly evolved flavour, which would have soothed his savage stomach ailments. He would have been like:

Damn, Emma! Holy s___ this is great apple-cake; here, try some! There is grandeur in this tasty cake, with its several flavours, having been originally cooked into a few baking trays or into one; and that, whilst this pub has gone on serving fine food according to the fixed hygiene laws of Tring, from so simple a beginning endless foods most beautiful and most wonderful have been, and are being, devoured.” And Emma, cake then firmly in hand, would have said something like, “My dear Charles, I shall try this enticing dessert, and I am glad to see you so enthused about something other than barnacles. Write a letter to Huxley or Lyell about that cake later. You need to focus on concocting an ending to that big species book of yours, not cakes. It’s been 20 bloody years, dude; cake can wait. End the book on a high note.” And so it must have happened.

Working at a museum collection is like having an extra home/office for a day or more. You get familiar with the environment while working there, and start to settle in and enjoy the local environs while taking work breaks. Or I do, anyway. So this post is also partly about how cake and other provisions are an important part, or even a perk, of life as a visiting museum researcher. Put in some good solid work, then it’s cake time, but where are the cakes? You explore, and you discover them– opening the door of an unfamiliar shop or pub near a museum can be like opening a museum cabinet to discover the goodness inside. Just don’t get them mixed up. Museum specimens: for research; subjects for science. Cakes: for eating; fuel for scientists. Got it?

But I digest digress. This post is not about my lunch. Not so much, anyway, although I did enjoy the cake quite a bit. Back to the chickens. Here, try some!

Darwins-chickens (1)Darwins-chickens (6)

Darwins-chickens (5) Darwins-chickens (4)

Above: Views of Darwin’s chickens laid out at the NHM-Tring.  (all photos in this post can be clucked to emchicken them)

The chickens, much like the pub lunch, did not disappoint in the least. Here I had before me Darwin’s own personal specimens, which I envisioned him dissecting and defleshing himself, studying them in deep introspection, then handing them over to the museum for curation once his lengthy researches were complete (all the ones I studied dated back to around 1863-1868, so they were curated shortly after The Origin was published (1859)). Perhaps the museum gave him some fine sponge-cake in return. There was at least one male and female adult of each of numerous breeds, many of them still bearing the dried flesh of centuries past. This was great for me, as the patella often gets removed and clucked chucked in the bin with its tendon when museum specimens of birds are prepared (much as elephant “sixth toe” sesamoids are). All of the specimens had their honking huge patellae on display, so that’s what a lot of my photos feature. I do so lament that I did not take a photo of the cake. Did I tell you about that cake? Oh… Check out these examples of Darwin’s chickens:

XXXX breed in right side view, with the patella indicated by a red arrow. It is still attached to the tibiotarsus by the patellar tendon (often misnamed the patellar "ligament", but it is just a continuation of the proximal tendon).

African rooster (wild variety? Darwin’s label was not clear) in right side view, with the patella indicated by a red arrow. That patella is still attached to the tibiotarsus by the patellar tendon (often misnamed the patellar “ligament”, but it is just a continuation of the proximal tendon).

Darwin's handwritten label and the well-endowed patella of the Spanish Cock. What? Oh, you. Stop it.

Darwin’s handwritten label and the well-endowed patella of the Spanish Cock. What? Oh, you. Stop it. That has nothing to do with cake, and only cake-related humour is allowed in this post.

Some other fascinating features exhibited by Darwin’s chickens, which he doubtless mulled over while nibbling on fine cakes, included the following:

The hindlimb of a Polish Silver Laced breed, nicely showing the ossified tendons (red arrow) along the tarsometatarsus. Why these tendons turn into bone is one of the great unsolved mysteries of bone biology/mechanics and avian evolution. Check out the famed feather crest here.

The hindlimb of a Polish Silver Laced breed, nicely showing the ossified tendons (red arrow) along the tarsometatarsus. Why these tendons turn into bone is one of the great unsolved mysteries of bone biology/mechanics and avian evolution.

Check out the famed feather crest of the Silver (Laced) Polish here; it gets so extreme in males that they have a hard time seeing, and can get beaten up by other cockerels when kept in mixed breed flocks.

Here on this blog, and of course on the companion blog “Towards the Chicken of the Future,” domestic chickens and wild junglefowl have often come up, most recently with the Dorking Chicken (another of Darwin’s own specimens that I studied) in the “Mystery Museum Specimen” poetry round of late. Dorkings are HUGE chickens; easily twice the weight of even a broiler chicken, up to 4-5kg. The Dorking-characteristic polydactyly featured in that post is also observed at a relatively high incidence in Silkie and Sultan breeds, I’ve learned. Like this one! (I was so patella-focused, or cake-somnolescent, that I missed it while studying at the museum and only noticed it now while browsing through my photos, bereft of cake)

Nice leg of a Sultan hen. There is an extra toe here as in the Dorking chicken; a duplicate hallux (first toe). This is not, as it might at first seem, a pathological condition as in modern "twisted toe"-suffering domestic chickens.

Nice leg of a Sultan hen. There is an extra toe here as in the Dorking chicken; a duplicate hallux (first toe). This is not, as it might at first seem, a pathological condition as in modern “twisted toe”-suffering domestic chickens.

Malays are another giant breed like the Dorking, but with longer and more muscular legs and longer necks, looking much more like a classic, badass wild junglefowl than a fancy, pampered chicken. But here, undressed to the bare bones, it just looks like a skinny chicken leg, albeit perhaps a bit svelte compared to the Dorking or Sultan.

Hindlimb of a Malay breed of chicken, which Wikipedia nicely tells the story of its misnomer (it may originate from Pakistan, not Malaysia!). Can you find the nice patella? Check out Darwin's lovely label, too.

Hindlimb of a Malay breed of chicken, which Wikipedia nicely tells the story of its misnomer (it may originate from Pakistan, not Malaysia!). Can you find the nice patella? Check out Darwin’s lovely label, too.

You may have come across wild-eyed news stories 5 years ago about “OMG Darwin was sooooooo wrong about chickens!”, referring to his writings on the origin of domestic chickens from Red junglefowl. As Greg Laden adeptly wrote, Darwin (say it with me) didn’t really get it very wrong after all. He did quite well, in fact. Some media outlets did get it more wrong, probably inspired by this press release. Oh well; the science they were reporting about definitely was interesting- modern chickens seem to have some of their yellow skin pigmentation-related genes from Grey junglefowl, although they are still largely descendants of Red junglefowl.

Here, have a JUMBLE-fowl, or rather a junglefowl cockerel, with another Darwin label:

Darwin's example of a wild-type chicken; a Red Junglefowl. As he suspected, these Asian birds were the ancestors of domestic chickens, but today evidence suggests that domestication occurred multiple times in Asia and with different wild varieties of junglefowl bred/mixed in different regions.

Darwin’s example of a wild-type chicken; a Red junglefowl. As he suspected, these Asian birds were the ancestors of domestic chickens, but today evidence indicates that domestication may have occurred multiple times in Asia and with different wild varieties of junglefowl bred/mixed in different regions.

Some breeds aren’t so funky inside, of course, but just have cool feather patterns on the outside, like the “pencilling” (dark streaks on white feathers) evident in pencil breeds; also called triple-laced. Like this fine chap below once would have had, before Darwin tore off his feathers and reduced him to a research-friendly naked skeleton:

A Golden Pencil Hamburgh breed of chicken (cockerel), whose skeleton features the leg and a fine articulated patella.

A Golden Pencil(led) Hamburg breed of chicken (cockerel), whose skeleton features the leg and a fine articulated patella.

Also known as the Holland Fowl, several European countries including the UK claim the Hamsburg as an original breed from their respective realm, and no surprise they do- it’s a lovely spangled chicken. Then, later in the 1800’s the Americans got involved in breeding them, too, and it’s all a big mess. They should get together, have some delectable cakes, and just sort it out.

Scaly, still-greasy foot and hindlimb of what Darwin labelled as the male of a "Game" breed.

Scaly, still-greasy foot and hindlimb of what Darwin labelled as the male of a “Game” breed.

We thus close with another leg of another chicken. Darwin was a bit naughty here, or else terminology of breeds has changed a lot since the 1850’s (very possible), as he just labelled this as a “Game” cockerel. Now, Gamefowl is a big category of breeds. I’m guessing this one was either (1) a Cornish/Indian Game variety or (2) an Old or Modern English Game Fowl. Maybe a person who knows their chicken breeding far better than me (that’s not hard!) will opine differently. The latter varieties were popular in Darwin’s time — the (Muffed) Old English version was mated with other breeds (Malay?) to produce the Modern English form as cockfighting “sports” became banned in 1849 and breeder attentions shifted to the polar opposite of producing showy, fancy birds instead. And thus the bufante, feathered-hair-adorned 1980s pop-rock group was created, to sing about mating or moulting or melting with people or something terribly disgusting and probably having nothing at all to do with chickens,  cake, or cockfighting, or other more seemly pursuits.

So, we have come to the end of my photos of Darwin’s chicken leg bones and such. If you’ve learned something here about chicken breeds, patellae, cake, or Darwin, that’s simply frabjous. Enough of those poncey pigeons, already! I’m crying fo… no, I won’t use that pun. Nevermind. Not even remotely cake-related. Let’s give Darwin’s chickens their just desserts, is the point– and a much better pun, too. Darwin’s chickens are an important part of Darwiniana, and an interesting evolutionary study in and of themselves. I’ve certainly become impressed during my researching for this blog post by the diverse, fascinating biology of chicken breeds. My copy of the “Complete Encyclopedia of Chickens” will be getting some more thorough reading shortly.

Today, however, I am off to return to the NHM-Tring and peruse their other, non-chickeny Galliformes and Anseriformes, with a detour to the mythical hoatzin. But… but… there may be a cake detour involved, too. I shall report back in due course. Off I go!

No, hopefully not that cake.

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Models of a basal dinosaur and bird, showing methods and key differences in body shape.

Our 3D computer models of a basal dinosaur and bird, showing methods and key differences in body shape. The numbers at the bottom are museum specimen numbers.

At about the moment I’m posting this, our Nature paper (our more formal page here, and the actual article here) embargo is ending, drawing a 14+ year gestation to a close. The paper is about how dinosaur 3D body shape changed during their evolution, and how that relates to changes in hindlimb posture from early dinosaurs/archosaurs to birds; “morpho-functional evolution” sums up the topic. We used the 3D whole-body computational modelling that I, Allen and Bates (among others) have developed to estimate evolutionary changes in body dimensions, rather than focusing on single specimens or (as in our last study) tyrannosaur ontogeny. We’ve strongly supported the notion (dating back to Gatesy’s seminal 1990 Paleobiology paper) that the centre of mass of dinosaurs shifted forwards during their evolution, and that this shift gradually led to the more crouched (flexed) hind leg posture that characterizes living birds. Here is a movie from our paper showing how we did the modelling:

And here is a summary of our 17 computer models of archosaur bodies, shown as one walks along the tips of the phylogeny shown in the video (the models are not considered to be ancestral to one another; we used a common computer algorithm called squared-change parsimony to estimate ancestral state changes of body dimensions between the 16 numbered nodes of the tree):

But we’ve done much more than just put numbers on conventional wisdom.

We’ve shown, to our own surprise, that the shift of the centre of mass was largely driven by evolutionary enlargements of the forelimbs (and the head and neck, and hindlimbs, to a less strong degree), not the tail as everyone including ourselves has assumed for almost 25 years. And the timing of this shift occurred inside the theropod dinosaur group that is called Maniraptora (or Maniraptoriformes, a slightly larger group), so the change began in animals that were not flying, but not long before flight evolved (depending on whom you ask, what taxonomy they favour and what evidence one accepts, either the smaller clade Eumaniraptora/Paraves or the bird clade Aves/Avialae).

Now, if you don’t like the cliche “rewriting the textbooks”, do have a look through texts on dinosaur/early avian palaeobiology and you probably will find a discussion of how the tail shortened, the centre of mass moved forwards as a consequence, the caudofemoral musculature diminished, and theropod dinosaurs (including birds) became more crouched as a result. We did that to confirm for ourselves that it’s a pretty well-accepted idea. Our study supports a large proportion of that idea’s reasoning, but modifies the emphasis to be on the forelimbs more than the tail for centre of mass effects, so the story gets more complex. The inference about caudofemoral muscles still seems quite sound, however, as is the general trend of increased limb crouching, but out paper approximates the timing of those changes.

Figure 3 from our paper, showing how the centre of mass moved forwards (up the y-axis) as one moves toward living birds (node 16); the funny dip at the end is an anomaly we discuss in the paper.

Figure 3 from our paper, showing how the centre of mass moved forwards (up the y-axis) as one moves toward living birds (node 16); the funny dip at the end is an anomaly we discuss in the paper.

A final implication of our study is that, because the forelimbs’ size influenced the centre of mass position, and thus influenced the ways the hindlimbs functioned, the forelimbs and hindlimbs are more coupled (via their effects on the centre of mass) than anyone has typically considered. Thus bipedalism and flight in theropods still have some functional coupling– although this is a matter of degree and not black/white, so by no means should we do away with helpful concepts like locomotor modules.

And in addition to doing science that we feel is good, we’ve gone the extra mile and presented all our data (yes, 17 dinosaurs’ worth of 3D whole body graphics!) and the critical software tools needed to replicate our analysis, in the Dryad database (link now working!), which should have now gone live with the paper! It was my first time using that database and it was incredibly easy (about 1 hour of work once we had all the final analysis’s files properly organized)– I strongly recommend others to try it out.

That’s my usual general summary of the paper, but that’s not what this blog article is about. I’ll provide my usual set of links to media coverage of the paper below, too. But the focus here is on the story behind the paper, to put a more personal spin on what it means to me (and my coauthors too). I’ll take a historical approach to explain how the paper evolved.


This paper’s story, with bits from the story of my life:

Embarassing picture of me before I became a scientist. Hardee's fast food restaurant cashier, my first "real job."

Embarassing picture of me before I became a scientist. Hardee’s fast food restaurant cashier, my first “real job”, from ~1999- no, wait, more like 1986. The 1980s-style feathered (and non-receding) hair gives it away!

Rewind to 1995. I started my PhD at Berkeley. I planned to use biomechanical methods and evidence to reconstruct how Tyrannosaurus rex moved, and started by synthesizing evidence on the anatomy and evolution of the hindlimb musculature in the whole archosaur group, with a focus on the lineage leading to Tyrannosaurus and to living birds. As my PhD project evolved, I became more interested and experienced in using 3D computational tools in biomechanics, which was my ultimate aim for T. rex.

In 1999, Don Henderson published his mathematical slicing approach to compute 3D body dimensions in extinct animals, which was a huge leap for the field forward beyond statistical estimates or physical toy models, because it represented dinosaurs-as-dinosaurs (not extrapolated reptiles/mammals/whatever) and gave you much more information than just body mass, with a lot of potential to do sensitivity analysis.

I struggled to upgrade my computer skills over the intervening years. I was developing the idea to reconstruct not only the biomechanics of T. rex, but also the evolutionary changes of biomechanics along the whole archosaur lineage to birds– because with a series of models of different species and a working phylogeny, you could do that. To me this was far more interesting than the morphology or function of any one taxon, BUT required you to be able to assess the latter. So Tyrannosaurus became a “case study” for me in how to reconstruct form and function in extinct animals, because it was interesting in its own right (mainly because of its giant size and bipedalism). (Much later, in 2007, I finally finished a collaboration to develop our own software package to do this 3D modelling, with Victor Ng-Thow-Hing and F. Clay Anderson at Honda and Stanford)

Me and a Mystery Scientist (then an undergrad; now a successful palaeontologist), measuring up a successful Cretaceous hypercarnivore at the UCMP; from my PhD days at Berkeley, ~2000 or so.

Me and a Mystery Scientist (then an undergrad; now a very successful palaeontologist!), measuring up a successful Cretaceous hypercarnivore at the UCMP; from my PhD days at Berkeley, ~2000 or so.

In all this research, I was inspired by not only my thesis committee and others at Berkeley, but also to a HUGE degree by Steve Gatesy, a very influential mentor and role model at Brown University. I owe a lot to him, and in a sense this paper is an homage to his trailblazing research; particularly his 1990 Paleobiology paper.

In 2001, I got the NSF bioinformatics postdoc I badly wanted, to go to the Neuromuscular Biomechanics lab at Stanford and learn the very latest 3D computational methods in biomechanics from Prof. Scott Delp’s team. This was a pivotal moment in my career; I became partly-an-engineer from that experience, and published some papers that I still look back fondly upon. Those papers, and many since (focused on validating and testing the accuracy/reliability of computer models of dinosaurs), set the stage for the present paper, which is one of the ones I’ve dreamed to do since the 1990s. So you may understand my excitement here…

Stanford's Neuromuscular Biomechanics Lab, just before I left in 2003.

Stanford’s Neuromuscular Biomechanics Lab, just before I left in 2003.

But the new paper is a team effort, and was driven by a very talented and fun then-PhD-student, now postdoc, Dr Vivian Allen. Viv’s PhD (2005-2009ish) was essentially intended to do all the things in biomechanics/evolution that I had run out of time/expertise to do in my PhD and postdoc, in regards to the evolution of dinosaur (especially theropod) locomotor biomechanics. And as I’d hoped, Viv put his own unique spin on the project, proving himself far better than me at writing software code and working with 3D graphics and biomechanical models. He’s now everything that I had hoped I’d become by the end of my postdoc, but didn’t really achieve, and more than that, too. So huge credit goes to Viv for this paper; it would never have happened without him.

We also got Karl Bates, another proven biomechanics/modelling expert, to contribute diverse ideas and data. Furthermore, Zhiheng Li (now at UT-Austin doing a PhD with Dr Julia Clarke) brought some awesome fossil birds (Pengornis and Yixianornis) from the IVPP in Beijing in order to microCT scan them in Londo. Zhiheng thus earned coauthorship on the paper — and I give big thanks to the Royal Society for funding this as an International Joint Project, with Dr Zhonghe Zhou at the IVPP.

That’s the team and the background, and I’ve already given you the punchlines for the paper; these are the primitive and the derived states of the paper. The rest of this post is about what happened behind the scenes. No huge drama or anything, but hard, cautious work and perseverance.

Me shortly after I moved to the RVC; video still frame from a dinosaur exhibit I was featured in. Embarassingly goofy pic, but I like the blurb at the bottom. It's all about the evolutionary polarity, baby!

Me shortly after I moved to the RVC; video still frame from a dinosaur exhibit (c. 2004) I was featured in. Embarassingly goofy pic, but I like the blurb at the bottom. It’s all about the evolutionary polarity, baby!

The paper of course got started during Viv’s PhD thesis; it was one of his chapters. However, back then it was just a focus on how the centre of mass changed, and the results for those simple patterns weren’t very different from those we present in the paper. We did spot, as our Nature supplementary information notes,  a strange trend in early theropods (like Dilophosaurus; to a lesser degree Coelophysis too) related to some unusual traits (e.g. a long torso) and suggested that there was a forward shift of centre of mass in these animals, but that wasn’t strongly upheld as we began to write the Nature paper.

On the urging of the PhD exam committee (and later the paper reviewers, too), Viv looked at the contributions of segment (i.e. head, neck, trunk, limbs, tail) mass and centre of mass to the overall whole body centre of mass. And I’m glad he did, since that uncovered the trend we did not expect to find: that the forelimb masses were far more important for moving the centre of mass forwards than the mass (or centre of mass) of the tail was– in other words, the statistical correlation of forelimb mass and centre of mass was strong, whereas changes of tail size didn’t correlate with the centre of mass nearly as much. We scrutinized those results quite carefully, even finding a very annoying bug in the 3D graphics files that required a major re-analysis during peer review (delaying the paper by ~6 months).

The paper was submitted to Nature first, passing a presubmission inquiry to check if the editor felt it fit the journal well enough. We had 3 anonymous peer reviewers; 1 gave extensive, detailed comments in the 3 rounds of review and was very fair and constructive, 1 gave helpful comments on writing style and other aspects of presentation as well as elements of the science, and 1 wasn’t that impressed by the paper’s novelty but wanted lots more species added, to investigate changes within different lineages of maniraptorans (e.g. therizinosaurs, oviraptorosaurs). That third reviewer only reviewed the paper for the first round (AFAIK), so I guess we won them over or else the editor overruled their concerns. We argued that 17 taxa were probably good enough to get the general evolutionary trends that we were after, and that number was ~16 more species than any prior studies had really done.

Above: CT scan reconstruction of the early extinct bird Yixianornis in slab conformation, and then Below: 3D skeletal reconstruction by Julia Molnar, missing just the final head (I find this very funny; Daffy Duck-esque) which we scaled to the fossil’s dimensions from the better data in our Archaeopteryx images. Yixianornis reconstruction There is also the concern, which the reviewers didn’t focus on but I could see other colleagues worrying about, that some of the specimens we used were either composites, sculpted, or otherwise not based on 100% complete, perfectly intact specimens. The latter are hard to come by for a diversity of extinct animals, especially in the maniraptoran/early bird group. We discussed some of these problems in our 3D Tyrannosaurus paper. The early dinosauromorph Marasuchus that we used was a cast/sculpted NHMUK specimen based on original material, as was our Coelophysis, Microraptor and Archaeopteryx; and our Carnegie ??Caenagnathus?? specimen was based more on measurements from 1 specimen than from direct scans, and there were a few other issues with our other specimens, all detailed in our paper’s Supplementary Information.

But our intuition, based on a lot of time spent with these models and the analysis of their data, is that these anatomical imperfections matter far, far less than the statistical methods that we employed– because we add a lot of flesh (like real animals have!) outside of the skeleton in our method, the precise morphology of the skeleton doesn’t matter much. It’s not like you need the kind of quality of anatomical detail that you need to do systematic analyses or osteological descriptive papers. The broad dimensions can matter, but those tend to be covered by the (overly, we suspect) broad error bars that our study had (see graph above). Hence while anyone could quibble ad infinitum about the accuracy of our skeletal data, I doubt it’s that bad– and it’s still a huge leap beyond previous studies, which did not present quantitative data, did not do comparative studies of multiple species, or did not construct models based on actual 3D skeletons as opposed to artists’ 2D shrinkwrapped reconstructions (the “Greg Paul method”). We also did directly measure the bodies of two extant archosaurs in our paper: a freshwater crocodile and a junglefowl (CT scan of the latter is reconstructed below in 3D).

One thing we still need to do, in future studies, is to look more carefully inside of the bird clade (Aves/Avialae) to see what’s going on there, especially as one moves closer to the crown group (modern birds). We represented modern birds with simply 1 bird: the “wild-type chicken” Red junglefowl, which isn’t drastically different in body shape from other basal modern birds such as a tinamou. Our paper was not about how diversity of body shape and centre of mass evolved within modern birds. But inspecting trends within Palaeognathae would be super interesting, because a lot of locomotor, size and body shape changes evolved therein; ostriches are probably a very, very poor proxy for the size and shape of the most recent common ancestor of all extant birds, for example, even though they seem to be fairly basal within that whole lineage. And, naturally, our study opens up opportunities for anyone to add feathers to our models and investigate aerodynamics, or to apply our methods to other dinosaur/vertebrate/metazoan groups. If the funding gods are kind to us, later this year we will be looking more closely, in particular, at the base of Archosauria and what was happening to locomotor mechanics in Triassic archosaurs…

Clickum to embiggum:

Australian freshwater crocodile, Crocodylus johnstoni; we CT scanned it in 3 pieces.

Australian freshwater crocodile, Crocodylus johnstoni; we CT scanned it in 3 pieces while visiting the Witmer lab in Ohio.

A Red junglefowl cockerel, spotted in Lampang, Thailand during one of my elephant gait research excursions there. Svelte, muscular and fast as hell.

A Red junglefowl cockerel, spotted in Lampang, Thailand during one of my elephant gait research excursions there. Svelte, muscular and fast as hell. This photo is here to remind me to TAKE BLOODY PICTURES OF MY ACTUAL RESEARCH SPECIMENS SO I CAN SHOW THEM!

I’d bore you with the statistical intricacies of the paper, but that’s not very fun and it’s not the style of this blog, which is not called “What’s in John’s Software Code?”. Viv really worked his butt off to get the stats right, and we did many rounds of revisions and checking together, in addition to consultations with statistics experts. So I feel we did a good job. See the paper if that kind of thing floats your boat. Someone could find a flaw or alternative method, and if that changed our major conclusions that would be a bummer– but that’s science. We took the plunge and put all of our data online, as noted above, so anyone can do that, and that optimizes the reproducibility of science.

What I hope people do, in particular, is to use the 3D graphics of our paper’s 17 specimen-based archosaur bodies for other things– new and original research, video games, animations, whatever. It has been very satisfying to finally, from fairly early in the paper-writing process onwards, present all of the complex data in an analysis like this so someone else can use it. My past modelling papers have not done this, but I aim to backtrack and bring them up to snuff like this. We couldn’t publish open access in Nature, but we achieved reasonably open data at least, and to me that’s as important. I am really excited at a personal level, and intrigued from a professional standpoint, to see how our data and tools get used. We’ll be posting refinements of our (Matlab software-based) tools, which we’re still finding ways to enhance, as we proceed with future research.

Velociraptor-model-min Dilophosaurus-model-min00

Above: Two of the 17 archosaur 3D models (the skinny “mininal” models; shrinkwrapped for your protection) that you can download and examine and do stuff with! Dilophosaurus on the left; Velociraptor on the right. Maybe you can use these to make a Jurassic Park 4 film that is better, or at least more scientifically accurate, than Hollywood’s version! ;-) Just download free software like Meshlab, drop the OBJ files in and go!

Now, to bring the story full circle, the paper is out at last! A 4 year journey from Viv’s PhD thesis to the journal, and for me a ~14 year journey from my mind’s eye to realization. Phew! The real fun begins now, as we see how the paper is received! I hope you like it, and if you work in this area I hope you like the big dataset that comes with it, too. Perhaps more than any other paper I’ve written, because of the long voyage this paper has taken, it has a special place in my heart. I’m proud of it and the work our team did together to produce it. Now it is also yours. And all 3200ish words of this lengthy blog post are, as well!

Last but not least, enjoy the wonderful digital painting that Luis Rey did for this paper (another of my team’s many failed attempts to get on the cover of a journal!); he has now blogged about it, too!

Dinosaur posture and body shape evolving up the evolutionary tree, with example taxa depicted.

Dinosaur posture and body shape evolving up the evolutionary tree, with example taxa depicted. By Luis Rey.


News stories about this paper will be added below as they come out, featuring our favourites:

1) NERC’s Planet Earth, by Harriet Jarlett: “Dinosaur body shape changed the way birds stand

2) Ed Yong on Phenomena: “Crouching  bird, hidden dinosaur

3) Charles Choi on Live Science: “Crouching bird, hidden evolutionary purpose?

4) Brian Handwerk on Nat Geo Daily News: “Birds’ “Crouching” Gait Born in Dinosaur Ancestors

5) Jennifer Viegas on Discovery News: “Heavier dino arms led evolution to birds

6) Puneet Kollipara on Science News: “Birds may have had to crouch before they could fly

And some superb videos- we’re really happy with these:

1) Nature’s “Crouching Turkey, Hidden Dragon

2) Reuters TV’s “3D study shows forelimb enlargement key to evolution of dinosaurs into birds

Synopsis: Decent coverage, but negligible coverage in the general press; just science-specialist media, more or less. I think the story was judged to be too complex/esoteric for the general public. You’d think dinosaurs, evolution, computers plus physics would be an “easy sell” but it was not, and I don’t think we made any big errors “selling” it. Interesting– I continue to learn more about how unpredictable the media can be.

Regardless, the paper has had a great response from scientist colleagues/science afficionados, which was the target audience anyway. I’m very pleased with it, too– it’s one of my team’s best papers in my ~18 year career.

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frontcover

The Unfeathered Bird book by Katrina van Grouw proclaims immediately in its Introduction that it “is not an anatomy of birds.”  True– it is far more than that, and it would be a shame if it had just been a dry, technical avian osteology reference book. It is a unique blend of art and science- particularly avian anatomy, evolution, taxonomy, natural history and more. The Unfeathered Bird is written for a general audience; birders/twitchers or just natural history buffs would be ideal targets of its unfettered passion for all things avian. A 12-year-old who is very keen on animals could enjoy it, and it may ignite the flames of ornithological excitement in many young or older readers. I am glad it was not called “The Naked Bird” as that would have caused some serious misconceptions (badum-tish!). The book is dripping with illustrations (at least one every two pages, often more). Almost all of the illustrations (except some paintings in the style of the cover) are in the same brownish sketch style that, like much of the book, evokes a bygone era of dark wooden cabinets and shadowed halls packed with skeletons, with nary an interactive graphics display, animatronic dinosaur or hyperdetailed cladogram in sight. It feels like an homage to the Victorian naturalists’ joy for anatomical detail conveyed through painstakingly detailed woodcuts. And while many still think of feathers as “the defining feature of birds,” enough about feathers already. Seriously. This is a book is about what lies beneath, and how all that non-fluffy stuff is important for birds’ lives, too.

(image-intense post; all can be clicked to embiggritate)

Katrina with peacock feather headdress? (back cover pic and rear view of same skeleton)

Katrina with peacock feather headdress? (back cover pic and rear view of same skeleton)

Katrina with front cover framed pic and the peacock skeleton that went with it.

Katrina with front cover framed pic and the peacock skeleton that went with it.

The Introduction continues to explain that the book is truly about how the external anatomy of birds is linked to the bony anatomy, which might remind astute readers of modern approaches like the extant phylogenetic bracket. The rest of the book uses both skeletal and unfeathered, quasi-myological illustrations to get this point across vividly. The explanatory text is written at a basic enough level for the average reader and is just the right length, with interesting anecdotes and natural history facts that even the expert reader will find interesting or even inspirational (e.g. possibly a goldmine for research ideas). First there is a 26 page “Basic” section with an introduction to avian osteology, with bountiful sketches to illustrate key organs and text explaining how it all fits together in the fully accoutered bird. The decision to use classical Linnean taxonomy (defunct or re-arranged taxa from the Systema Naturae like Accipitres, Picae, Anseres, Grallae, Gallinae and Passeres; which are the six “Specific” chapters in the second section of the book) was a good one- it enhances the classical feel of the tome and gives the author a great opportunity to discuss convergent evolution and how that misled past ornithologists.

But for me, the book is most pleasurable for the visualizations and the passion for all things birdy that weaves through them and the accompanying text. The removal of feathers, or even all soft tissues, from bird bodies (posed in naturalistic behaviours) that van Grouw renders in her illustrations shows birds in a new light, emphasizing the strangeness and diversity that lie beneath. The author begins the book with a touching Acknowledgments section in which her husband Hein van Grouw, curator of birds at the Natural History Museum’s Tring collection, features very prominently, making it clear that the book was a team operation and comes from the heart after a 25-year journey. This gives the book a special warmth that is preserved throughout the remainder- although the illustrations are of flayed bodies or boiled / beetle-macerated skeletons, the tone is nothing less than an earnest love for birds of all kinds, and a zest for portraying those feelings to the reader in sketches and prose. It is a joyous celebration, not a somber litany, of the wonder of birds that can be gleaned from dead bodies. There is so much powerful, awesome imagery stuffed into those pages that it is hard to summarize. I’ll let five of my favourite images from the book (more are in her gallery and her book’s Facebook page; but even these are just the tip of the icebird) help get this across (used with permission of the author):

Naked kiwi in action.

Naked kiwi in action.

The unscaled bird: guineafowl feet.

The unscaled bird: guineafowl feet.

Deplumed sparrowhawk with dove trophy, exalting in its triumph.

Deplumed sparrowhawk with dove trophy, exalting in its triumph.

Budgerigar has made a friend? Or came to grips with its own mortality?

Budgerigar has made a friend? Or came to grips with its own mortality?

Trumpet Manucode WTF anatomy! Spiraling tracheal coil made me gasp in awe when I saw this image in the book.

Trumpet Manucode’s WTF anatomy! Spiraling tracheal coil made me gasp in awe when I saw this image in the book.

Now I’ll depart from this post just being a book review. I went to the Tring collection to do some research, and arranged my trip so I’d also get to see the debut of a Tring special exhibit featuring The Unfeathered Bird, and also to meet Katrina as well as Hein van Grouw. The placement of the exhibit at Tring is apropos, because Katrina was a curator at the museum until a few years ago and Hein still is. But the inspiration for the work and the specimens used (with a few exceptions, including from other museums) are Katrina’s. She (with Hein’s help) procured bodies of birds to dissect, macerate and sketch for the book over its 25 year fledging period, noting in the Acknowledgments that “no birds were harmed” to do this– do read those acknowledgments, as there are some amusing tales there of how she obtained some specimens.

I was fortunate to be able to take some photos of the exhibit while they set it up, and grabbed some candid images of Katrina and colleagues during that process. The following images show off the exhibit, which is all in one clean, bright, simply adorned room in the Tring that lets Katrina’s framed sketches be the focus. Here are some examples:

Poster advert for the book in the Tring collections.

Poster advert for the book in the Tring collections.

Tring exhibit setup, with Katrina, husband Hein, and helper finishing it up.

Tring exhibit setup, with Katrina, husband Hein, and helper finishing it up.

Tring exhibit now ready.

Tring exhibit now ready.

Tring exhibit case.

Tring exhibit case.

Framed sketches at Tring exhibit.

Framed sketches at Tring exhibit.

Framed sketches at Tring exhibit.

More framed sketches at Tring exhibit.

The exhibit is fun for people who are already Unfeathered Bird fans, and a good way of drawing in new ones. The book is a precious thing that any fan of birds, especially scientists, really needs to have a hard copy of. While it claims not to be an anatomy text, its illustrations provide ample opportunities to use it for that purpose. But really the point of owning all 287-plus pages is to bask in the warmth of true, pure appreciation for classic ornithology, which I found infectious. It is a book by and for bird lovers, but also for those that find the interface of art and science to be fascinating.

I confess I used to hate birds. I found them annoying and boring; all that flitting and twitting and pretentious feathers. “Get over yourselves, already, and calm down too!” was my reaction to them. When I started grad school, I had an open disdain for birds, even moreso than for mammals (OK, except cats). I was a “herp” fan through and through, for most of my life (childhood spent catching anoles in Florida, or stalking frogs in Ohio; during visits to my grandparents). What won me over was studying birds (and eventually mammals, too) as a young scientist, and learning how incredible they are– not just as endpoints in the story of theropod dinosaur evolution, as my thesis focused on, but as amazing animals with spectacular form-function relationships.  The Unfeathered Bird is saturated with that amazement, so we’re birds of an unfeather.

Framed sketch of dodo head at Tring exhibit.

Framed sketch of dodo head at Tring exhibit.

Entirely unfeathered Indian peafowl in matching views.

Entirely unfeathered Indian peafowl in matching views.

Painted Stork and Toco Toucan sketches.

Painted Stork and Great Hornbill sketches.

Red junglefowl, wild ancestor of domestic chickens (and the book ends with several such breeds illustrated),

Red junglefowl, wild ancestor of domestic chickens (and the book ends with several such breeds illustrated).

Katrina told me that she is already deep into writing the next book, whose subject I won’t spoil for you here but maybe we will be lucky enough to have her appear in the Comments and plug it? :) (Her website does say “It was Hein’s stroke of genius to include domestic birds and they’ve provided the inspiration for my next project.” so the cat is out of the bag and amongst the pigeons!) It is great to hear that the book has done quite well sales-wise and critically, such as ~#67 on the Amazon sales list at one point– I hope this paves the way for more such books not only from Katrina, but from others engaged in lateral thinking (and still others) on the boundaries of science-art.

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…a daily picture of anatomy! And today it is three pictures, scoob-a-dee!

Welcome back again to Freezermas! 

For the previous days of Freezermas we first had 1 picture, then 2, now guess how many we have today? Right, we’ve settled into a groove and have three (plus one silly one). Today is fresh beefy anatomy day! No focus on bones, but on soft tissues– however, once again, I’m representin’ bird legs! And this time, no mystery things to identify; sorry. But if you want to muscle in on some myology, today is the day for you. I will unwrap the thigh of an ostrich and consider the major muscles that power rapid running in this biped, and how they illuminate the evolution of bipedal motion along the line of descent to birds. For more ostrich escapades, see this old post. And we’re off!

Stomach-Churning Rating: 7/10; plenty of fresh, red, meaty meat from ostrich leg muscles.

Ostrich thigh muscles 1

Here you are looking at a right hindlimb of an ostrich, in side/lateral view. To help orient yourself, the hip lies deep in the middle of the image and the knee is the rounded bump near the bottom right corner, with the shank angling sharply back toward the bottom left.

I’ve labelled six muscles in yellow. As usual for sauropsid (bird/reptile) pelvic limb muscles, they have sensible names that reflect their attachments. They don’t have so many silly old mammalian names like pectineus or latissimus, which tell you rather little about the muscles themselves. We can thank 19th century anatomists like two of my anatomist heroes, Hans Gadow and Alfred Romer (who refined Gadow’s earlier work and made it more popular among English-speakers and palaeontologists), for that enlightened nomenclature.

The six muscles seen above are the IC (iliotibialis cranialis), IL (iliotibialis lateralis), “AMB2?” (one of the ambiens muscles– correctly identified; ignore the ?), ITC (iliotrochantericus caudalis), CFP (caudofemoralis pars pelvica) and FCLP (a mouthful to say: flexor cruris lateralis pars pelvica). The ambiens is the one oddly, non-anatomically named muscle, and has nothing to do with helping you sleep (pssst– wake up! Muscles are exciting!), but everything to do with the state of total awesomeness, which is what “ambiens” means. Maybe. Or I am making shit up.

Amazed ostrich

The IC, IL and AMB2 are parts of the triceps femoris group (discussed in my 1st Freeezermas post), or for mammal fans the quadriceps. The IC and AMB are in front of the hip so they flex it (move the thigh forward; protract it); the IL is right around the hip so it can flex or extend the hip (protract/retract the femur); all three of these can extend (straighten) the knee joint to varying degrees. The IC is fairly typical for a bird except for its size, and helps to quickly swing the leg through the air between steps. Some birds have multiple parts of the IL, but ostriches and many others have simplified it to one major mass; regardless, it is a major muscle used to support the weight of the body.

The AMB2 is a remarkable muscle unique to ostriches; it can also be called the dorsal ambiens muscle. Typical birds just have a single head of the AMB sitting on the preacetabular (pubic) tubercle, so in front of and below the hip. It has a crazy tendon that snakes past the knee (in some birds, perforating/grooving the patella) into the lower leg muscles and may be able to even pull on the toes. But ostriches, for some reason, added a second head of this muscle that was shifted way up onto the front of the pelvis (the ilium; dorsal bladelike bone). Crocodilians also have a 2nd ambiens muscle but in a different position, and almost certainly as an example of convergent evolution. The function of the ambiens is mysterious, but this muscle has featured prominently in avian systematics/taxonomy, evolution (invoked as a key muscle used in perching) and more.

These muscles of the triceps femoris group are easily identifiable in crocodiles and other reptiles because they are remarkably similar in their attachments. The main changes these muscles experienced during the evolution of bipedalism, dinosaurs and later birds are simply proportional– they got bigger, with stronger, larger attachments on the pelvis and the front of the knee (the CC/LC, if you remember from Freezermas day 1).

The ITC is a muscle that is very dear to me. I’ve written a lot about it, and I love saying the name “Iliotrochantericus caudalis”- it is musical to me. For mammal fans, think gluteal muscles (medial gluteal in particular). It is a huge, pennate muscle (short and strongly angled muscle fibres in a “sandwich” with a tendinous sheet between the two layers of fibres). It has a short, broad tendon that wraps around the trochanteric crest (a structure on the upper front end of the femur with a history that goes wayyyy back into dinosaurs; long story!) to insert in a scarred depression. The ITC seems to mainly rotate the femur around its long axis to help support the body. I could go on and on about this muscle, which is part of the enigmatic “deep dorsal” thigh muscle group — the homologies of this group among land vertebrates are still controversial and confusing. But I will spare you the on-and-on. Incidentally, the ITC  is the “oyster” in birds that is the best cut of meat. And in ostriches it makes a massive steak.

The CFP also has a cool evolutionary history. It runs from the back of the pelvis to the middle of the femur, closely adjoined to the caudal head of the muscle (CFC), which is more vestigial. In birds the CFP is usually not a large muscle, but in other sauropsids/reptiles it can be fairly hefty, although almost never as hefty as its more famous counterpart the caudofemoralis longus (= CFC in birds). Probably any dinosaur specialist is familiar with its origin and its insertion: respectively, the “brevis fossa” on the back of the ilium; a big shelf of bone; and the fourth trochanter of the femur; a crest of bone that is reduced to a scar/tubercle in birds. Much like its tail-based counterpart, the CFP became progressively reduced closer and closer to birds. This is related to a reduction in the amount of movement of the femur/thigh during locomotion, as birds shortened their tails and shifted their balance forward, as Steve Gatesy showed in a classic 1990 paper. Hopefully there will be more about this subject in a future paper from my team…

The FCLP is another muscle that didn’t change much, except by getting larger as we trace its evolution from early reptiles to birds. It is a “hamstring” muscle that is an important power source during locomotion in birds like the ostrich, because it retracts the lower limb (flexes the knee; hence flexor cruris in its name) as well as the femur/thigh (extends the hip). Your semitendinosus muscle is a good comparison to it. Indeed, these two differently named muscles are homologous– our very distant tetrapod ancestor had the same single muscle, and its descendants didn’t change it that much on our lineage or on the avian/reptile one.

Ostrich thigh muscles 3

I’ve reflected the IL muscle out of the way so we can see the second layer of muscles underneath it. Now we see two more muscles of the thigh, and large ones at that– the FMTL (femorotibialis lateralis) and ILFB (iliofibularis).

The FMTL simply is a part of the triceps femoris group that only comes from the femur and hence only, but due to its large size powerfully, straightens the knee. Unlike the other muscles in this group, it has no action about the hip joint. It is very similar to your vastus lateralis muscle: its fleshy origin dominates the surface of the femur (thigh bone). There are two other parts of that muscle, hidden in this figure, much like our vastus group has multiple parts. Again, this is a muscle that enlarged on the lineage leading to modern birds.

And that evolutionary enlargement applies, too, to the ILFB, whose prominent insertion I discussed on day 1 of Freezermas. This huge “biceps” muscle (it is single-headed unlike in humans, so the name “biceps” does not apply well) is the most powerful of the “hamstring”-type muscles that extend the hip and flex the knee. Therefore it is important for the “knee-driven” locomotion of birds. And hence the ILFB enlarged during avian evolution– which is very evident from changes of both its bony origin on the back of the pelvis/ilium and its insertion on the fibula.

Ostrich thigh muscles 2

Here, for the terminus of today’s trio of struthious tributes and tribulations, I’ve moved the ILFB  out of the way so you can see the various inner/medial layer of thigh muscles. Some of the former muscles are more exposed now, and we can see three new ones: the FCM (flexor cruris medialis), PIFM+(PIF)L (the tongue-twisting puboischiofemoralis medialis et lateralis), and tiny ISF (ischiofemoralis).

The FCM (~mammalian semitendinosus) is merely another, smaller part of the FCLP’s “hamstring” group, and its thin tendon blends with that of the FCLP, so it very much works with that muscle in locomotion, and has a similar evolutionary history.

The PIFM+L are “adductors”, but in birds they don’t really do any adduction (drawing the legs inwards) because they are right behind, rather than below or inside, the hip. They act as hip extensors/retractors of the femur, and probably aid more in holding the femur steady (“postural muscles”) than playing a major role in producing power for locomotion like the ILFB/hamstring group does. In earlier reptiles, they were much more important, for preventing the legs from splaying too far away from the body.

The ISF is usually quite a large muscle in birds, but ostriches and some other ratites have reduced it to a thin slip of muscle– often mistaken for other muscles (indeed, like a few other muscles I’ve described here, modern anatomists still get confused by this muscle– an otherwise superb recent description by Gangl et al., among others, mis-identifies this and some other muscles— an error an upcoming paper from my group will rectify). Normally the ISF sits atop a bone-free window on the outer surface of the pelvis, the ilio-ischiadic fenestra (literally a window in Latin) in birds; in ostriches it has moved more onto the ischium. In contrast, in other sauropsids it lies inside the pelvis, so during its evolution it became more lateral, but the insertion on the upper femur was maintained. It is a weak rotator and extensor of the hip, especially in ostriches in which its role is probably proportionately puny.

And there you have read a healthy chunk of my 2001 PhD thesis, condensed into less jargonious language. You might now know almost half of the key muscles of the avian hind limb. If you made it this far, you are one awesome anatomical enthusiast. If you eat meat, apply this lesson to the next chicken thigh you consume, to consumate this enthusiasm.

A broader point I’d like to make here is that anatomy is best conveyed not only along with the functional narrative (How does anatomy work?) but also the evolutionary tale (Where did anatomy come from and What were the consequences of its changes? Why did it change?). This takes it away from dry memorization of terms and locations, and carries it into the realm of explaining why nature is the way it is, and how every organism’s biology has a richly detailed and complex background. This style portrays nature as much more like that tangled bank that Darwin so enchantingly envisioned. I’ve tried to do that justice here, using this one ostrich whom we affectionately called Twinkletoes, or Twinkie, when we dissected it back in 2002.

Happy Freezermas! Sing it: “On the third day of Freezermas, this blo-og gave to me: one tibiotarsus, two silly pictures, a-and three muscle layers from Twinkie!”

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