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Posts Tagged ‘tetrapod’

Who needs “Ice Road Truckers” when you have the “John’s Freezer” team on the road with fossils, amphibians, felids and 3D phenotype fun? No one, that’s who. We’re rocking the Cheltenham Science Festival for our first time (as a group), and pulling out all the stops by presenting two events! Here’s the skinny on them, with updates as the week proceeds.

Stomach-Churning Rating: 2/10 for now (just bones), but it could change once the cheetah dissection is under way… 8/10 bloody cheetah bits but only at the end (updated)

Right now, Lauren Sumner-Rooney (of “Anatomy To You” and other fame) is on-site with a rotating team of others from our lab, in the “Free Activity Tents” area of the Imperial Gardens/Square, inside a marquee where we’ll be showing off our NERC-funded tetrapod research all week. This “First Steps” event features not only our past and present work with Jenny Clack, Stephanie Pierce, Julia Molnar and others on Ichthyostega & its “fishapod” mates, but also our “scampering salamanders” research in Spain, Germany and England. I’ve blogged a lot about all that, and won’t repeat it here, but you can see a 3D-printed Ichthyostega skeleton, view the skeleton in a virtual reality 3D environment, see related specimens and engage in kid-friendly activities, and talk to our team about this and other related research.

Ichthyostega 3D printed backbone is born!

Ichthyostega 3D printed backbone is born!

The central themes of that event are how bone structure relates to function and how we can use such information, along with experimental measurements and computer models of real salamanders, to reconstruct how extinct animals might have behaved as well as how swimming animals became walking ones. How did fins transform into limbs and what did that mean for how vertebrates made the evolutionary transition onto land? If you know my team’s work, that encapsulates our general approach to many other problems in evolutionary biomechanics (e.g. how did avian bipedalism evolve?). Added benefits are that you too can explore this theme in a hands-on way, and you can talk with us about it in person. That continues all week (i.e. until Saturday evening); I’ll be around from Thursday afternoon onwards, too. Kids of all ages are welcome!

Ichthyostega 3D print taking shape!

Ichthyostega 3D print taking shape!

Then, on Saturday for our second free event we join forces with Ben Garrod (master of primate evolution, the secrets of bones, and “Attenborough and the Giant Dinosaur”) and RVC’s forensic pathologist Alexander Stoll as well as Sophie Regnault (“sesamoid street” PhD student w/me). As the “Large Animal Dissection” title hints, it’s not the right kind of gig to bring small kids to. There will be blood and stuff— we’ll be dissecting a cheetah together from 10am-4pm. This will involve walking through all the major organ systems, giving evolutionary anecdotes, and plenty more, with an aim to understand how the magnificent adaptations of cheetahs evolved—but also to investigate what problem(s) this animal faced that led to its sad demise. By the day’s end, there will just be a skeleton left. Get a front row seat early for this event, which serendipitously ties into “Team Cat”’s Leverhulme Trust-funded research project (we wanted a big animal and it just happened to be a cheetah; I had hoped for a giant croc or a shark or something but can’t complain!).

Ichthyostega 3D print is ready!

Ichthyostega 3D print is ready!

If you miss these events, please do cry bitter tears of regret. But don’t despair, there will be another “big cat dissection” in the London area in ~November (watch here for details), and plenty more fossil tetrapod stuff to come, plus a LOT more dinosaurs on the horizon!

Guess the bones! (photo by Zoe Self)

Guess the bones! (photo by Zoe Self)

And please come back to this blog post for more pics and stories as the week carries on… For hashtag afficionados, you can follow the fun on Twitter etc. at #firststepsCSF16. What a world we live in!

Update 1: While you’re here, check out our Youtube playlists of tetrapod-related videos:

Lobe-finned fishes

Ichthyostega‘s awesome anatomy

Tetrapod evolution: Tiktaalik to salamanders!

Update 2: Photos of our main stand (about tetrapod evolution)

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Our poster/banner display looks nice.

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Our tent brings in some punters.

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Our bones excite people young and old, sighted and blind.

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Fun with stickers and lab t-shirts.

And…

Update 3: Cheetah meat & greet
Ben, Alex, Sophie and I tackled the cheetah dissection today and it went GREAT! Much better than I’d optimistically expected. Rain didn’t scare the crowds off and neither did the gore, which there was some of (gelatinous spinal cords, lumpy tumors and at least one flying tiny bit of cheetah flesh that landed on a good-natured audience member!). Photos will tell the tale:

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Peek-a-boo!

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Sophie and Alex help us get set up in our tent.

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Our initial rough schedule- although we ended up improvising more after lunch.

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Dissectors assemble!

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The beast revealed. It was skinned by the museum that loaned it to us.

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Alex showing his talent: removing the viscera in one piece from end to end, starting with the tongue.

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Impressive finding of a surgical fixture (plate and wires) on the tibia, which had been used to hold the shattered bone back together long enough for it to heal. Added to the kidney disease and liver-spleen-lung cancer, this cheetah was in the sorriest shape of any cadaver I’ve seen yet.

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Cheetah coming to pieces: (from bottom) lumbar/pelvic region, hindlimb, thorax, forelimb and other bits.

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Dr Adam Rutherford, an eye expert, did a nice dissection of the cheetah’s eye, here showing the tapetum lucidum (reflective membrane), which shows up as light blue colour. Its small size befits the not-very-nocturnal habits of cheetahs.

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The lens of the cheetah’s eye. Now cloudy because of dehydration and crystalization, but still fascinating to see.

Want to see more images and the enthusiastic responses from the audience (we got some great feedback)? Check out Twitter’s #cheltscifest feed, or more simply my Storify condensation of the cheetah-related tweets here.

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Happy Darwin Day from the frozen tundra sunny but muddy, frosty lands of England! I bring you limb muscles as peace offerings on this auspicious day. Lots of limb muscles. And a new theme for future blog posts to follow up on: starting off my “Better Know A Muscle” (nod to Stephen Colbert; alternative link) series. My BKAM series intends to walk through the evolutionary history of the coolest (skeletal/striated) muscles. Chuck Darwin would not enjoy the inevitable blood in this photo-tour, but hopefully he’d like the evolution. Off we go, in search of better knowledge via an evolutionary perspective!

There is, inarguably, no cooler muscle than M. caudofemoralis longus, or CFL for short. It includes the largest limb muscles of any land animal, and it’s a strange muscle that confused anatomists for many years– was it a muscle of the body (an axial or “extrinsic” limb muscle, directly related to the segmented vertebral column) or of the limbs (an “abaxial” muscle, developing with the other limb muscles from specific regions of the paraxial mesoderm/myotome, not branching off from the axial muscles)? Developmental biologists and anatomists answered that conclusively over the past century: the CFL is a limb muscle, not some muscle that lost its way from the vertebral column and ended up stranded on the hindlimb.

The CFL is also a muscle that we know a fair amount about in terms of its fossil record and function, as you may know if you’re a dinosaur fan, and as I will quickly review later. We know enough about it that we can even dare to speculate if organisms on other planets would have it. Well, sort of…

Stomach-Churning Rating: 8/10. Lots of meaty, bloody, gooey goodness, on and on, for numerous species. This is an anatomy post for those with an appetite for raw morphology.

Let’s start from a strong (and non-gooey) vantage point, to which we shall return. The CFL in crocodiles and most other groups is (and long was) a large muscle extending from much of the front half or so of the tail to the back of the femur (thigh bone), as shown here:

Julia Molnar's fabulous illustration of Alligator's limb muscles, from our 2014 paper in Journal of Anatomy.

Julia Molnar’s fabulous illustration of Alligator‘s limb muscles, from our 2014 paper in Journal of Anatomy. Note the CFL in blue at the bottom right.

As the drawing shows, the CFL has a friend: the CFB. The CFB is a shorter, stumpier version of the CFL restricted to the tail’s base, near the hip. The “B” in its name means “brevis”, or runty. It gets much less respect than its friend the CFL. Pity the poor CFB.

But look closer at the CFL in the drawing above and you’ll see a thin blue tendon extending past the knee to the outer side of the lower leg. This is the famed(?) “tendon of Sutton“, or secondary tendon of the CFL. So the CFL has two insertions, one on the femur and one (indirectly) onto the shank. More about that later.

Together, we can talk about these two muscles (CFL and CFB) as the caudofemoralis (CF) group, and the name is nice because it describes how they run from the tail (“caudo”) to the femur (“femoralis”). Mammal anatomists were late to this party and gave mammal muscles stupidly unhelpful names like “gluteus” or “vastus” or “babalooey”. Thanks.

But enough abstract drawings, even if they rock, and enough nomenclature. Here is the whopping big CFL muscle of a real crocodile:

Huge Nile crocodile, but a relatively small CFL.

Huge Nile crocodile, but a relatively small CFL.

Bigger crocs have smaller legs and muscles.

Bigger crocs have smaller legs and thus smaller leg muscles, relatively speaking. CFL at the top, curving to the left.

The giant Nile croc's CFL muscle removed for measurements.

The giant Nile croc’s CFL muscle removed for measurements. 2.35 kg of muscle! Not shabby for a 278 kg animal.

However, maybe crocodile and other archosaur CFL muscles are not “average” for leggy vertebrates? We can’t tell unless we take an evolutionary tack to the question.

Where did the CFL come from, you may ask? Ahh, that is shrouded in the fin-limb transition‘s mysteries. Living amphibians such as salamanders have at least one CF muscle, so a clear predecessor to the CFL (and maybe CFB) was present before reptiles scampered onto the scene.

But going further back through the CF muscles’ history, into lobe-finned fish, becomes very hard because those fish (today) have so few fin muscles that, in our distant fishy ancestors, would have given rise eventually to the CF and other muscle groups. With many land animals having 30+ hindlimb muscles, and fish having 2-8 or so, there obviously was an increase in the number of muscles as limbs evolved from fins. And because a limb has to do lots of difficult three-dimensional things on land while coping with gravity, more muscles to enable that complex control surely were needed.

OK, so there were CF muscles early in tetrapod history, presumably, anchored on that big, round fleshy tail that they evolved from their thin, finned fishy one — but what happened next? Lizards give us some clues, and their CFL muscles aren’t all that different from crocodiles, so the CFL’s massive size and secondary “tendon of Sutton” seems to be a reptile thing, at least.

Courtesy of Emma Schachner, a large varanid lizard's very freshly preserved CFL and other hindlimb muscles.

Courtesy of Emma Schachner, a large varanid lizard’s very freshly preserved CFL and other hindlimb muscles.

Courtesy of Emma Schachner, zoomed in on the tendons and insertions of the CFL muscle and others.

Courtesy of Emma Schachner, zoomed in on the tendons and insertions of the CFL muscle and others. Beautiful anatomy there!

Looking up at the belly of a basilisk lizard and its dissected right leg, with the end of the CFL labelled.

Looking up at the belly of a basilisk lizard and its dissected right leg, with the end of the CFL labelled. It’s not ideally dissected here, but it is present.

An unspecified iguanid(?) lizard, probably a juvenile Iguana iguana, dissected and showing its CFL muscle at its end. The muscle would extemd about halfway down the tail, though.

An unspecified iguanid(?) lizard, probably a juvenile Iguana iguana, dissected to reveal its CFL muscle near its attachment to the femur. The muscle would extend further, about halfway down the tail, though.

Let’s return to crocodiles, for one because they are so flippin’ cool, and for another because they give a segue into archosaurs, especially dinosaurs, and thence birds:

A moderate-sized (45kg) Nile crocodile with its CFL muscle proudly displayed.

A moderate-sized (45kg) Nile crocodile with its CFL muscle proudly displayed. Note the healthy sheath of fat (cut here) around the CFL.

American alligator's CFL dominates the photo. Photo by Vivian Allen.

American alligator’s CFL dominates the photo [by Vivian Allen].

Black caiman, Melanosuchus, showing off its CFL muscle (pink "steak" in the middle of the tail near the leg).

Black caiman, Melanosuchus, showing off its CFL muscle (pink “steak” in the middle of the tail near the leg), underneath all that dark armour and fatty superficial musculature.

A closer look at the black caiman's thigh and CFL muscle.

A closer look at the black caiman’s thigh and CFL muscle.

Like I hinted above, crocodiles (and the anatomy of the CFL they share with lizards and some other tetrapods) open a window into the evolution of unusual tail-to-thigh muscles and locomotor behaviours in tetrapod vertebrates.

Thanks in large part to Steve Gatesy’s groundbreaking work in the 1990s on the CFL muscle, we understand now how it works in living reptiles like crocodiles. It mainly serves to retract the femur (extend the hip joint), drawing the leg backwards. This also helps support the weight of the animal while the foot is on the ground, and power the animal forwards. So we call the CFL a “stance phase muscle”, referring to how it mainly plays a role during ground contact and resisting gravity, rather than swinging the leg forwards (protracting the limb; i.e. as a “swing phase muscle”).

The “tendon of Sutton” probably helps to begin retracting the shank once the thigh has moved forward enough, facilitating the switch from stance to swing phase, but someone really needs to study that question more someday.

And thanks again to that same body of work by Gatesy (and some others too), we also understand how the CFL’s anatomy relates to the underlying anatomy of the skeleton. There is a large space for the CFL to originate from on the bottom of the tail vertebrae, and a honking big crest (the fourth trochanter) on the femur in most reptiles that serves as the major attachment point, from which the thin “tendon of Sutton” extends down past the knee.

Femur bones (left side) from an adult ostrich (Left) and Nile crocodile (Right).

Femur bones (left side; rear view) from an adult ostrich (left) and Nile crocodile (right). Appropriate scale bar is appropriate. The fourth trochanter for the CFL is visible in the crocodile almost midway down the femur. Little is left of it in the ostrich but there is a bumpy little muscle scar in almost the same region as the fourth trochanter, and this is where the same muscle (often called the CFC; but it is basically just a small CFL) attaches.

That relationship of the CFL’s muscular anatomy and the underlying skeleton’s anatomy helps us a lot! Now we can begin to look at extinct relatives of crocodiles; members of the archosaur group that includes dinosaurs (which today we consider to include birds, too), and things get even more interesting! The “tendon of Sutton”, hinted at by a “pendant” part of the fourth trochanter that points down toward the knee, seems to go away multiple times within dinosaurs. Bye bye! Then plenty more happens:

A large duckbill dinosaur's left leg, with a red line drawn in showing roughly where the CFL would be running, to end up at the fourth trochanter. Many Mesozoic dinosaurs have skeletal anatomy that indicates a similar CFL muscle.

A large duckbill dinosaur’s left leg, with a red line drawn in showing roughly where the CFL would be running, to end up at the fourth trochanter. Many Mesozoic dinosaurs have skeletal anatomy that indicates a similar CFL muscle.

We can even go so far as to reconstruct the 3D anatomy of the CFL in a dinosaur such as T. rex ("Sue" specimen here; from Julia Molnar's awesome illustration in our 2011 paper), with a fair degree of confidence.

We can even go so far as to reconstruct the 3D anatomy of the CFL in a dinosaur such as T. rex (“Sue” specimen here; from Julia Molnar’s awesome illustration as part of our 2011 paper), with a fair degree of confidence. >180kg steak, anyone?

As we approach birds along the dinosaur lineage, the tail gets smaller and so does the fourth trochanter and thus so must the CFL muscle, until we’re left with just a little flap of muscle, at best. In concert, the hindlimbs get more crouched, the forelimbs get larger, flight evolves and voila! An explosion of modern bird species!

Ozburt (72)

Left femur of an ostrich in side view (hip is toward the right side) showing many muscles that attach around the knee (on the left), then the thin strap of CF muscle (barely visible; 2nd from the right) clinging near the midshaft of the femur.

Another adult ostrich's CF muscle complex, removed for study.

Another adult ostrich’s CF muscle complex, removed for study. Not enough ostrich myology for you yet? Plenty more in this old post! Or this one! Or this one… hey maybe I need to write less about ostriches? The CF muscle complex looks beefy but it’s no bigger than any other of the main hindlimb muscles, unlike the CFL in a crocodile or lizard, which puts everything else to shame!

STILL not enough ostrich for you yet? Take a tour of the major hindlimb muscles in this video:

And check out the limited mobility of the hip joint/femur here. No need for much femur motion when you’re not using your hip muscles as much to drive you forwards:

But I must move on… to the remainder of avian diversity! In just a few photos… Although the CF muscles are lost in numerous bird species, they tend to hang around and just remain a long, thin, unprepossessing muscle:

Chicken's right leg in side view. CFC (equivalent of CFL) muscle outlined and labelled.

Chicken’s right leg in side view. CFC muscle (equivalent of CFL; the ancestral CFB is confusingly called the CFP in birds, as it entirely resides on the pelvis) outlined and labelled.

A jay (species?) dissected to show some of the major leg muscles, including the CF. Photo by Vivian Allen.

A jay (species? I forget) dissected to show some of the major leg muscles, including the CFL-equivalent muscle; again, smallish. [Photo by Vivian Allen]

Finally, what’s up with mammals‘ tail-to-thigh CF-y muscles? Not much. Again, as in birds: smaller tail and/or femur, smaller CF muscles. Mammals instead depend more on their hamstring and gluteal muscles to support and propel themselves forward.

But many mammals do still have something that is either called the M. caudofemoralis or is likely the same thing, albeit almost always fairly modest in size. This evolutionary reduction of the CF muscle along the mammal (synapsid) lineage hasn’t gotten nearly as much attention as that given to the dinosaur/bird lineage’s CFL. Somebody should give it a thoroughly modern phylogenetic what-for! Science the shit outta that caudofemoralis…

Yet, oddly, to give one apparent counter-example, cats (felids) have, probably secondarily, beefed up their CF muscle a bit:

Cats have a pretty large CF muscle in general, and this jaguar is no exception! But mammals still tend to have fairly wimpy tails and thus CF muscles, or they even lose them (e.g. us?).

Cats have a pretty large CF muscle in general, and this jaguar is no exception! But mammals still tend to have fairly wimpy tails and thus CF muscles, or they even lose them (e.g. us?). [photo by Andrew Cuff, I think]

In summary, here’s what happened (click to embeefen):

Better Know A Muscle: The Evolution of M. caudofemoralis (longus)

Better Know A Muscle: the evolution of M. caudofemoralis (longus).

I hope you enjoyed the first BKAM episode!
I am willing to hear requests for future ones… M. pectoralis (major/profundus) is a serious contender.

P.S. It was Freezermas this week! I forgot to mention that. But this post counts as my Freezermas post for 2016; it’s all I can manage. Old Freezermas posts are here.

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[This is the original, unedited text of my shorter, tighter (and I think actually better) News & Views piece for Nature, on the paper described below)

Ambitious experimental and morphological studies of a modern fish show how a flexible phenotype may have helped early “fishapods” to make the long transition from finned aquatic animals into tetrapods able to walk on land.

Stomach-Churning Rating: 1/10. Cute fish. Good science. Happy stomachs!

Photo by Antoine Morin, showing Polypterus on land.

Photo by Antoine Morin, showing Polypterus on land.

Napoleon Bonaparte’s military excursions into Egypt in 1798-1799 led a young French naturalist, Ètienne Geoffroy Saint-Hilaire, to cross paths with a strange fish that had paired lungs and could “walk” across land on its stubby, lobelike fins. In 1802, he dubbed this fish “Polyptère bichir”1, today known as the Nile bichir, Polypterus bichir La Cepède 1803. The bichir’s mélange of primitive and advanced traits helped to catapult Geoffroy into scholarly conflict with the reigning naturalist Georges Cuvier back in France and to establish Ètienne as a leading anatomist, embryologist and early evolutionary researcher of repute even today2. Now, on their own excursion under the very “evo-devo” flag that the discoverer of Polypterus helped raise, Canadian scientists Standen et al.3 suggest how the remarkable plasticity of the skeleton of Polypterus (the smaller west African relative of P. bichir, P. senegalus or “Cuvier’s bichir”) reveals a key part of the mechanism that might have facilitated the gradual transition from water to land and thus from “fishapods” to tetrapods (four-limbed vertebrates).

In a bold experiment, the authors raised 149 young bichirs on land and in water for eight months, then studied how they moved on land vs. in water, and also how the ultimate shape of the skeletal elements of the paired front fin bases differed between the land- and water-raised bichirs. Standen et al.3 discovered that both the form and function of the fins’ foundations transformed to better satisfy the constraints of moving on land. Land-acclimated bichirs took faster steps on land, their fins slipped across the substrate less, they held their fins closer to their body, their noses stayed more aloft and their tails undulated less, with less variable motions overall—behaviours that the authors had predicted should appear to enhance walking abilities on land. In turn, the bones of the neck and shoulder region altered their shape to produce a more mobile fin base with greater independence of fin from neck motion, along with improved bracing of the ventral “collarbone” region. These environmentally-induced traits should have fostered the locomotor changes observed in “terrestrialized” fish and aided the animals in resisting gravity, and they represent a common biological phenomenon termed developmental plasticity4,5. Interestingly, the land-reared fish could still swim about as well as the wholly aquatic cohort, so there was not a clear trade-off between being a good swimmer and a good walker, which is surprising.

Considered alone, the developmental plasticity of bichir form and function shows how impressive these amphibious fish are. But Standen et al.’s study3  ventured further, to apply the lessons learned from bichir ontogeny to a phylogenetic context and macroevolutionary question. The phenotypic plasticity during bichir development, they infer, could have been harnessed during the evolutionary transformation of fins for swimming into limbs for walking, in the “fishapod” ancestors of tetrapods. Indeed, bichirs are close to the base of the family tree of fishes6, and other living relatives of tetrapods have reduced or lost their fins (lungfishes) or adapted to strange deep-sea swimming lifestyles, never walking on land (coelacanths). Thus perhaps bichirs and the “fishapod” lineage share what Geoffroy would have called “unity of type”, today termed homology, of their developmental plasticity in response to a land environment. Surveying the fossil record of early “fishapods” and tetrapods, Standen et al.3 found that the macroevolutionary changes of neck and shoulder anatomy in these gradually more land-adapted animals parallel those they observed in terrestrialized Polypterus, providing ancillary support for their hypothesis.

A further test of the application of Polypterus’s plasticity to fossil tetrapods is naturally difficult. However, the “fishapod” lineage has some exceptional examples of fossil preservation. With sufficient sample sizes (e.g. fossil beds that reveal growth series, such as the Late Devonian Miguasha site in Canada7) and palaeoenvironmental gradients in fish or tetrapods, one could imagine performing a rigorous indirect test. Even small samples could be helpful– for example, the early tetrapod Ichthyostega exhibits some developmental changes in its forelimb suggesting that it became more terrestrial as it grew, whereas the related Acanthostega does not evidence such changes8— this hints at some developmental plasticity in the former animal.

During the Devonian period (~360-420 million years ago), were the “fishapod” ancestors of tetrapods floundering about on land now and then, gradually shifting from anatomy and behaviours that were more developmentally plastic (as in bichirs) to ones that were more canalized into the terrestrialized forms and functions that more land-adapted tetrapods retained? An attractive possibility is that the developmental plasticity could have led to fixation (reduction of plasticity), an evolutionary phenomenon called genetic assimilation, which another intellectual descendant of Geoffroy, Conrad Hal Waddington, promoted from the 1950s onwards9, a concept that now enjoys numerous cases of empirical support10 that this one may eventually join.

The nature of the genetic and developmental mechanism that bichirs use to achieve the observed developmental plasticity is still unclear. If it has a high enough degree of heritability, then it could be selected for in cross-generational experiments with bichirs. With sufficient time and luck raising these unusual fish, the hypothesis that their plastic response to a terrestrial environment can become genetically assimilated could be directly tested. This study could thus become an epic exemplar of how genetic assimilation can contribute not only to microevolutionary change but also to major macroevolutionary events, as was presciently suggested in a seminal review of developmental plasticity4.

This genetic assimilation is the Polypterus study’s reasonable speculation, and one that Geoffroy likely would have applauded, all the more for involving his beloved bichirs. Much as Napoleon’s landfall in Egypt was not a lasting success, bichirs never left wholly terrestrial descendants despite their malleable locomotor system. But the same type of plastic developmental mechanism that bichirs use today to make tentative, floppy incursions of the terrestrial realm might have been harnessed by our own “fishapod” forebears, leaving a far more revolutionary dynasty upon the Earth.

 

References

  1.  Geoffroy, E. (1802). Histoire naturelle et description anatomique d’un nouveau genre de poisson du Nil, nommé polyptère. Annales du Muséum d’Histoire Naturelle 1:57-68.
  2. Le Guyader, H., & Grene, M. (2004) Geoffroy Saint-Hilaire: A Visionary Naturalist. Univ. Chicago Press.
  3. Standen, E. M., Du, T. Y., & Larsson, H. C. E. (2014). Developmental plasticity and the origin of tetrapods. Nature, published online.
  4. West-Eberhard, M. J. (1989). Phenotypic plasticity and the origins of diversity. Annual Review of Ecology and Systematics 20:249-278.
  5. Pigliucci, M., Murren, C. J., & Schlichting, C. D. (2006). Phenotypic plasticity and evolution by genetic assimilation. Journal of Experimental Biology 209(12):2362-2367.
  6. Near, T. J., Dornburg, A., Tokita, M., Suzuki, D., Brandley, M. C., & Friedman, M. (2014). Boom and bust: ancient and recent diversification in bichirs (Polypteridae: Actinopterygii), a relictual lineage of ray‐finned fishes. Evolution 68:1014-1026.
  7. Cloutier, R. (2013). Great Canadian Lagerstätten 4. The Devonian Miguasha Biota (Québec): UNESCO World Heritage Site and a Time Capsule in the Early History of Vertebrates.Geoscience Canada40:149-163.
  8. Callier, V., Clack, J. A., & Ahlberg, P. E. (2009). Contrasting developmental trajectories in the earliest known tetrapod forelimbs.Science324:364-367.
  9. Waddington, C. H. (1953). Genetic assimilation of an acquired character. Evolution 7:118-126.
  10. Crispo, E. (2007). The Baldwin effect and genetic assimilation: revisiting two mechanisms of evolutionary change mediated by phenotypic plasticity. Evolution 61:2469-2479.

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Construction of the Phyletisches Museum in Jena, Germany began on Goethe’s birthday on August 28, 1907. The Art Nouveau-styled museum was devised by the great evolutionary biologist, embryologist and artist/howthefuckdoyousummarizehowcoolhewas Ernst Haeckel, who by that time had earned fame in many areas of research (and art), including coining the terms ontogeny (the pattern of development of an organism during its lifetime) and phylogeny (the pattern of evolution of lineages of organisms through time) which feature prominently in the building’s design and exhibits (notice them intertwined in the tree motif below, on the front of the museum). Ontogeny and phylogeny, and the flamboyant artistic sensibility that Haeckel’s work exuded, persist as themes in the museum exhibits themselves. Haeckel also came up with other popular words such as Darwinism and ecology, stem cell, and so on… yeah the dude kept busy.

Cavorting frogs from Haeckel's masterpiece Kunstformen der Natur (1904).

Cavorting frogs from Haeckel’s masterpiece Kunstformen der Natur (1904).

I first visited the Phyletisches Museum about 10 years ago, then again this August. Here are the sights from my latest visit: a whirlwind ~20 minute tour of the museum before we had to drive off to far-flung Wetzlar. All images are click-tastic for embiggenness.

Stomach-Churning Rating: 3/10 for some preserved specimens. And art nouveau.

Willkommen!

Willkommen!

Frog ontogeny, illustrated with gorgeous handmade ?resin? models.

Frog ontogeny, illustrated with gorgeous handmade ?resin? models.

Fish phylogeny, illustrated with lovely artistry.

Phylogeny of Deuterostomia (various wormy things, echinoderms, fish and us), illustrated with lovely artistry.

Phylogeny of fish and tetrapods.

Phylogeny of fish and tetrapods.

Slice of fossil fish diversity.

Slice of fossil fish diversity.

Plenty of chondryichthyan jaws and bodies.

Plenty of chondrichthyan jaws/chondrocrania, teeth and bodies.

Awesome model of a Gulper eel (Saccopharyngiformes).

Awesome model of a Gulper Eel — or, evocatively, “Sackmaul” auf Deutsch (Saccopharyngiformes).

Lobe-finned fishes (Sarcopterygii)- great assortment.

Lobe-finned fishes (Sarcopterygii)- great assortment including a fossil coelacanth.

Lungfish body/model and skeleton.

Lungfish body and skeleton.

Coelacanth!

Coelacanth!

Coelacanth staredown!

Coelacanth staredown!

Fire salamander! We love em, and the museum had several on display- given that we were studying them with x-rays, seeing the skeleton and body together here in this nice display was a pleasant surprise.

On into tetrapods– a Fire Salamander (Salamandra salamandra)! We love ’em, and the museum had several on display- given that we were studying them with x-rays, seeing the skeleton and body together here in this nice display was a pleasant surprise.

A tortoise shell and skeleton, with a goofball inspecting it.

A tortoise shell and skeleton, with a goofball inspecting it.

In a subtle nod to recurrent themes in evolution, the streamlined bodies of an ichthyosaur and cetacean shown in the main stairwell of the museum, illustrating convergent evolution to swimming locomotor adaptations.

In a subtle nod to recurrent themes in evolution, the streamlined bodies of an ichthyosaur and cetacean shown in the main stairwell of the museum, illustrating convergent evolution to swimming adaptations.

Phylogeny of reptiles, including archosaurs (crocs+birds).

Phylogeny of reptiles, including archosaurs (crocs+birds).

Gnarly model of an Archaeopteryx looks over a cast of the Berlin specimen, and a fellow archosaur (crocodile).

Gnarly model of an Archaeopteryx looks over a cast of the Berlin specimen, and a fellow archosaur (crocodile). The only extinct dinosaur on exhibit!

Kiwi considers the differences in modern bird palates: palaeognathous like it and fellow ratites/tinamous (left), and neognathous like most living birds.

Kiwi considers the differences in modern bird palates: palaeognathous like it and fellow ratites/tinamous (left), and neognathous like most living birds.

Echidna skeleton. I can't get enough of these!

Echidna skeleton. I can’t get enough of these!

Skulls of dugong (above) and manatee (below).

Skulls of dugong (above) and manatee (below), Sirenia (seacows) closely related to elephants.

Fetal manatee. Awww.

Fetal manatee. Awww.

Adult Caribbean manatee, showing thoracic dissection.

Adult Caribbean manatee, showing thoracic dissection.

Hyraxes, which Prof. Martin Fischer, longtime curator of the Phyletisches Museum, has studied for many years.  Rodent-like elephant relatives.

Hyraxes, which Prof. Martin Fischer, longtime curator of the Phyletisches Museum, has studied for many years. Rodent-like elephant cousins.

Old exhibit at the Phyletisches Museum, now gone: Forelimbs of an elephant posed in the same postures actually measured in African elephants, for the instant of foot touchdown (left pic) and liftoff (right pic). Involving data that we published in 2008!

Old exhibit at the Phyletisches Museum, now gone: Forelimbs of an elephant posed in the same postures actually measured in African elephants, for the instant of foot touchdown (left pic) and liftoff (right pic). Involving data that we published in 2008!

Gorilla see, gorilla do. Notice "bent hip, bent knee" vs. "upright modern human" hindlimb postures in the two non-skeletal hominids.

Eek, primates! Gorilla see, gorilla do. Notice the primitive “bent hip, bent knee” vs. the advanced “upright modern human” hindlimb postures in the two non-skeletal hominids.

Phylogeny of select mammals, including the hippo-whale clade.

Phylogeny of artiodactyl (even-toed) mammals, including the hippo-whale clade.

Hand (manus) of the early stem-whale Ambulocetus.

Hand (manus) of the early stem-whale Ambulocetus.

Carved shoulderblade (scapula) of a bowhead whale (Balaena mysticetus), which apparently Goethe owned. Quite a relic!

Carved shoulderblade (scapula) of a bowhead whale (Balaena mysticetus), which apparently Goethe owned (click to emwhalen and read the fine print). Quite a relic!

One of Haeckel's residences. There is also a well-preserved house of his that one can visit, but I didn't make it there.

One of Haeckel’s residences, across the street from the museum. There is also a well-preserved house of his that one can visit, but I didn’t make it there. I heard it’s pretty cool.

Jena is tucked away in a valley in former East Germany, with no local airport for easy access- but get to Leipzig and take a 1.25 hour train ride and you’re there. Worth a trip! This is where not just ontogeny and phylogeny were “born”, but also morphology as a modern, rigorous discipline. Huge respect is due to Jena, and to Haeckel, whose quotable quotes and influential research still resonate today, in science as well as in art.

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Short and sweet post here; it’s sunny outside and I want to be there BBQing!

I had a buried folder of CT files labelled as a species of fish, but on digging them out and segmenting them I realize it is not what I expected (inner fish or not!), as you will see.

Stomach-Churning Rating: 2/10; simple CT scan of a body.

Mystery Anatomy 2014same rules as before; remember that the scoreboard has been reset.

Identify the animal in the CT scout/pilot image below, as specifically as you can. But… (READ THE SENTENCE BELOW FIRST BEFORE ANSWERING!)

Today’s special rule: Summertime is coming and that means superhero films! Your answer must be in the form of a dialogue between a superhero(ine) and a supervillain(ess)! 

Difficulty: Even I am not 100% sure what this is but I have a decent idea. Not super hard, but not a super good segmentation.

Pow! Bam! Biff! Go forth and conquer! Then invite the Human Torch to your BBQ.

 

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Jason Anderson, vertebrate paleontologist and anatomist at the University of Calgary (Canada), shared these two intriguing photos with me, and agreed for me to share them with you. Yay, thanks Jason! Good timing for a badly needed Mystery Dissection post:

What are they (species ID) *AND* from what region of the body?

(they are the same region, same specimen, same animal)

RULE: Your answer must be in the form of a winter holiday song (at least four lines)!

If you’ve been following some of my recent tweets, I’ve been tweeting about the “joys” of increased academic paperwork around holiday-time; e.g. this one:

As always, you can score extra points for creativity.

Remember: the scoreboard is here.

Difficulty: Integumentary

Stomach-Churning Rating: 6/10; if beauty is only skin deep, then ugly is, too?

Here they are…

(more…)

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I love doing sciencey road trips with my team when I can. Last week, we got a treat: four of us got a behind-the-scenes tour of the fairly new Crocodiles of the World facility near Oxford; just over 90 minutes west of our lab, nestled in the pictureseque Cotswolds region. We were not disappointed, so you get to share in the joy! In photo-blog format. Pics can be clicked to emcrocken.

In the midst of an unpreposessing industrial estate lies: AWESOME!

In the midst of an unpreposessing industrial estate lies: AWESOME!

If you want to bone up on your croc species, go here and here. I won’t go into details. This is an eye candy post!

Reasonably accurate description that caught my eye.

Reasonably accurate description that caught my eye. My scientific interest in crocodiles starts here, and with their anatomy/relationship with dinosaurs, but I’ve loved crocs since I was an infant (one of my first words, as I may have written here before, was “dock-a-dile”, for my favourite stuffed animal at the time [R.I.P.]).

Siamese crocodiles. They were apart when we entered, then got snuggly later, as I've often seen this species do.

Siamese crocodiles. The large male is “Hugo.” They were apart when we entered, then got snuggly later, as I’ve often seen this species do. Heavily endangered (<300 in the wild?), so any breeding is a good thing!

The above photo brings me to one of my general points. Crocodiles of the World seems genuinely to be a centre that is breeding crocodiles for conservation purposes (and for education, entertainment and other zoo-like stuff). Essentially every crocodile enclosure had a mated pair, and several were breeding. Such as…

Yes, that is a Dwarf African crocodile, Osteolaemus, and it is a female on her nest-mound. Which means...

Yes, that is a Dwarf African crocodile, Osteolaemus, and indeed it is a female on her nest-mound. Which means…

Eggs of said Osteolaemus.

Eggs of said Osteolaemus.

And babies of said Osteolaemus!

And babies of said Osteolaemus! As if the adults aren’t cute enough with their short snouts and doglike size/appearance! These guys have striking yellow colouration, too. I’d never seen it in person before.

That’s not all!

Male American Alligator warming up. Smaller female partner lives in same enclosure.

Male American Alligator “Albert” warming up. Smaller female partner “Daisy” lives in same enclosure. Plenty of babies from these guys, too! Daisy comes when called by name, and Albert is learning to do so.

~1 meter long juvenile Nile crocodiles, bred at the facility.

~1 meter long juvenile Nile crocodiles, bred at the facility.

But then crocodile morphological diversity (colours, textures) and behaviour is just too cool not to focus on a bit, so here are some highlights from our visit!

Endearing shot of a crocodylian I seldom get to see anywhere: Paleosuchus trigonatus, the Cuvier's Dwarf Caiman. Spiny armoured hide and quite terrestrial; poorly known in many ways. Some more info is here- http://crocodilian.com/paleosuchus/description.html (note its tortured taxonomy)

Endearing shot of a crocodylian I seldom get to see anywhere: Paleosuchus trigonatus, the Schneider’s Dwarf Caiman. Spiny armoured hide and quite terrestrial; poorly known in many ways. Some more info is here (note its tortured taxonomy)

Black caiman, Melanosuchus niger, showing some interest in us.

Black caiman, Melanosuchus niger, showing some interest in us.

Cuban crocodiles cooling off by exposing their mouths.

Cuban crocodiles (Crocodylus rhombifer; pound for pound the most badass croc in my experience; badassitude that this photo captures nicely) cooling off by exposing the well-vascularized soft tissues of the mouth region.

But it’s not just crocs there, either, and some of the highlights were non-croc surprises and memorable encounters:

A surprisingly friendly and tame Water monitor (14 yrs old; does kids parties). Note person for scale.

A surprisingly friendly and tame Water monitor (14 yrs old; does kids parties). Note person for scale. Was about 2 meters long, 20 kg or so.

Business end of nice Water monitor, with tongue engaged.

Business end of nice Water monitor, with tongue engaged.

And we got a nice farewell from an African spur-thigh tortoise (Geochelone sulcata) with an oral fixation (action sequence thereof):

tortoise-nom (1)
tortoise-nom (2)

tortoise-nom (3)
tortoise-nom (4)

tortoise-nom (5)

Chowmp!

If someone visits this facility and leaves without being converted to a croc-lover, they must be from a different planet than me. It is a celebration of crocodiles; the owner, Shaun Foggett, is the real deal. He sold his home and quit his job as a carpenter to care for crocodiles, and it seems to be a great success– about to get greater, as they have plans to move to a new, bigger, proper site! They are seeking funding, so if you can contribute go here.

Right then… UK residents and visitors: you need to go here! Badly! Get off the blog and go now. If it is a Saturday/Sunday (the cramped industrial estate location only allows the public then).

Otherwise just stew and imagine how much fun you could be having checking out crocodiles. I cruelly posted this on a Tuesday to ensure thorough marination of any croc-geeks.

Muhaha!  ;-)

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