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Let's play find-the-spandrel!

Let’s play find-the-spandrel!

We just passed the 35th anniversary of the publication of Gould and Lewontin’s classic, highly cited, highly controversial essay (diatribe?), “The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme.” The 21st of September 1979 was the fateful date. Every PhD student in biology should read it (you can find pdfs here– this post assumes some familiarity with it!) and wrestle with it and either love it or hate it- THERE CAN BE NO MIDDLE GROUND! With some 5405 citations according to Google Scholar, it has generated some discussion, to put it lightly. Evolutionary physiologists and behaviourists who were working at the time it came out have told me stories of how it sent (and continues to send) shockwaves through the community. Shockwaves of “oh crap I should have known better” and “Hell yeah man” and “F@$£ you Steve,” more or less.

I am among those who love “The Spandrels Paper“. I love it despite its many flaws that people have pointed out to seemingly no end- the inaccurate architectural spandrel analogy, the Gouldian discursive (overly parenthetical [I'm a recovering victim of reading too much Gould as an undergrad]) writing style, the perhaps excessive usage of “Look at some classic non-scientific literature I can quote”, the straw men and so on. I won’t belabour those; again your favourite literature search engine can be your guide through that dense bibliography of critiques. I love it because it is so daringly iconoclastic, and because I think it is still an accurate criticism of what a LOT of scientists who do research overlapping with evolutionary biology (that is, much of biology itself) do.

The aspects of The Spandrels Paper that I still think about the most are:

(1) scientists seldom test hypotheses of adaptation; they are quick to label something that is useful to an animal as an adaptation and then move on after rhapsodizing about how cool of an adaptation it is; and

(2) thus alternatives to adaptation, which might be very exciting topics to study in their own right, get less attention or none.

True for #2, evo-devo has flourished by raising the flag of constraint (genetic/developmental/other factors that prevent evolution from going in a certain direction, or even accelerate it in less random directions). That’s good, and there are other examples (genetic drift, we’ve heard about that sometimes), but option #1 still often tends to be the course researchers take. To some degree, labelling something as an adaptation is used as hype, to make it more exciting, I think, in plenty of instances.

Truth be told, much as Gould and Lewontin admitted in their 1979 paper and later ones, natural selection surely forges lineages that have loads of adaptations (even in the strictest sense of the word), and a lot of useful traits of organisms are thus indeed adaptations by any stripe. But the tendency seems to be to assume that this presumptive commonality of adaptations means that we are justified to quickly label traits as adaptations.

Or maybe some researchers just don’t care about rigorous tests of adaptation as they’re keen to do other things. Standards vary. What I wanted to raise in this post is how I tend to think about adaptation:

I think adaptations are totally cool products of evolution that we should be joyous to imagine, document, test and discover. But that means they should be Special. Precious. A cause for celebration, to carefully document by scientific criteria that something is an adaptation in the strictest sense, and not a plesiomorphy/exaptation (i.e. an adaptation at a different level in the evolutionary hierarchy; or an old one put to new uses), spandrel/byproduct, or other alternatives to adaptation-for-current-biological-role.

But that special-ness means testing a hypothesis of adaptation is hard. As many authors waving the flag of The Modern Comparative Method (TMCM) have contended, sciencing truth-to-adaptationist-power by the rules of TMCM takes a lot of work! George Lauder’s 1996 commentary in the great Adaptation book (pdf of the chapter here) outlined a lengthy procedure of  “The Argument from Design“; i.e., testing adaptation hypotheses. At its strictest implementation it could take a career (biomechanics experiments, field studies, fitness measurements, heritability studies, etc.) to test for one adaptation.

Who has time for all that?

The latter question seems maladaptive, placing cart and horse bass-ackwards. If one agrees that adaptations are Special, then one should be patient in testing them. Within the constraints of the practical, to some degree, and different fields would be forced to have different comfort levels of hypothesis testing (e.g. with fossils you can’t ever measure fitness or other components of adaptation directly; that does not mean that we cannot indirectly test for adaptations– with the vast time spans available, one would expect palaeo could do a very good job of it, actually!).

I find that, in my spheres of research, biomechanists in particular tend to be fast to call things they study adaptations, and plenty of palaeontologists do too. I feel like over-usage of the label “adaptation” cheapens the concept, making the discovery of one of the most revered and crucial concepts in all of evolutionary biology seem cheapened and trite. Things that are so easy to discover don’t seem as precious. When everything is awesome, nothing is…

I’ve always hesitated, thanks in part to The Spandrels Paper’s indoctrination, from calling features of animals adaptations, especially in my main research. I nominally do study major ?adaptations? such as terrestrial locomotion at giant body sizes, or the evolution of dinosaurian bipedalism. I searched through my ~80 serious scientific papers lately and found about 50 mentions of “adapt” in an adaptationist, evolutionary context. That’s not much considering how vital the concept is (or I think it is) to my research, but it’s still some mentions that slipped through, most of them cautiously considered– but plenty more times I very deliberately avoided using the term. So I’m no model of best practice, and perhaps I’m too wedded to semantics and pedantry on this issue, but I still find it interesting to think about, and I’ve gradually been headed in the direction of aspect #2 (above in bold) in my research, looking more and more for alternative hypotheses to adaptation that can be tested.

I like talking about The Spandrels Paper and I like some of the criticism of it- that’s healthy. It’s a fun paper to argue about and maybe we should move on, but I still come back to it and wonder how much of the resistance to its core points is truly scientific. I’m entering into teaching time, and I always teach my undergrads a few nuggets of The Spandrels Paper to get them thinking about what lies beyond adaptation in organismal design.

 What do other scientists think? What does adaptation mean (in terms of standards required to test it) to you? I’m curious how much personal/disciplinary standards vary. How much should they?

For the non-scientists, try this on for size: when our beloved Sir David Attenborough (or any science communicator) speaks in a nature documentary about how the otter is “perfectly adapted” to swim after prey underwater, do you buy into that or question it? Should you? (I get documentaries pushing me *all the time* to make statements like this, with a nudge and a wink when I resist) Aren’t scientists funny creatures anyway?

MysteryCT12
Here’s an image that struck me as cool and possibly perplexing. And so we have another Mystery Anatomy post! Brought to you by some free time on my current trip to Gondwanaland.

Stomach-Churning Rating: 1/10; simple CT scan slice… of something.

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

Identify the animal in the CT slice shown above, as specifically as you can. No special rules.

Difficulty: Plenty.

Begin!

 

I’ll let the poll (prior post) run for a while but as it winds down I wanted to explain why I posted it:

In the past, I’ve often run into scientists who, when defending their published or other research, respond something like this:

“Yeah those data (or methods) might be wrong but the conclusions are right regardless, so don’t worry.”

And I’ve said things like that before. However, I’ve since realized that this is a dangerous attitude, and in many contexts it is wrong.

If the data are guesses, as in the example I gave, then we might worry about them and want to improve them. The “data are guesses” context that I set the prior post in comes from Garland’s 1983 paper on the maximal speeds of mammals– you can download a pdf here if this link works (or Google it). Basically the analysis shows that, as mammals get bigger, they don’t speed up as a simple linear analysis might show you. Rather, at a moderate size of around 50-100kg body mass or so, they hit a plateau of maximal speed, then bigger mammals tend to move more slowly. However, all but a few of the data points in that paper are guesses, many coming from old literature. The elephant data points are excessively fast in the case of African elephants, and on a little blog-ish webpage from the early 2000s we chronicled the history of these data– it’s a fun read, I think. The most important, influential data plot from that paper by Garland is below, and I love it– this plot says a lot:

Garland1983

I’ve worried about the accuracy of those data points for a long time, especially as analyses keep re-using them– e.g. this paper, this one, and this one, by different authors. I’ve talked to several people about this paper over the past 20 years or so. The general feeling has been in agreement with Scientist 1 in the poll, or the quote above– it’s hard to imagine how the main conclusions of the paper would truly be wrong, despite the unavoidable flaws in the data. I’d agree with that statement still: I love that Garland paper after many years and many reads. It is a paper that is strongly related to hypotheses that my own research seeks out to test. I’ve also tried to fill in some real empirical data on maximal speeds for mammals (mainly elephants; others have been less attainable), to improve data that could be put into or compared with such an analysis. But it is very hard to get good data on even near-maximal speeds for most non-domesticated, non-trained species. So the situation seems to be tolerable. Not ideal, but tolerable. Since 1983, science seems to be moving slowly toward better understanding of the real-life patterns that the Garland paper first inferred, and that is good.

But…

My poll wasn’t really about that Garland paper. I could defend that paper- it makes the best of a tough situation, and it has stimulated a lot of research (197 citations according to Google; seems low actually, considering the influence I feel the paper has had).

I decided to do the poll because thinking about the Garland paper’s “(educated) guesses as data” led me to think of another context in which someone might say “Yeah those data might be wrong but the conclusions are right regardless, so don’t worry.” They might say it to defend their own work, such as to deflect concerns that the paper might be based on flawed data or methods that should be formally corrected. I’ve heard people say this a lot about their own work, and sometimes it might be defensible. But I think we should think harder about why we would say such things, and if we are justified in doing so.

We may not just be making the best of a tough situation in our own research. Yes, indeed, science is normally wrong to some degree. A more disconcerting situation is that our wrongs may be mistakes that others will proliferate in the future. Part of the reasoning for being strict stewards of our own data is this: It’s our responsibility as scientists to protect the integrity of the scientific record, particularly of our own published research because we may know that best. We’re not funded (by whatever source, unless we’re independently wealthy) just to further our own careers, although that’s important too, as we’re not robots. We’re funded to generate useful knowledge (including data) that others can use, for the benefit of the society/institution that funds us. All the more reason to share our critical data as we publish papers, but I won’t go off on that important tangent right now.

In the context described in the latter paragraph and the overly simplistic poll, I’d tend to favour data over conclusions, especially if forced to answer the question as phrased. The poll reveals that, like me, most (~58%) respondents also would tend to favour data over conclusions (yes, biased audience, perhaps- social media users might tend to be more savvy about data issues in science today? Small sample size, sure,  that too!). Whereas very few (~10%) would favour conclusions, in the context of the poll. The many excellent comments on the poll post reveal the trickier nuances behind the poll’s overly simplistic question, and why many (~32%) did not favour one answer over the other.

If you’ve followed this blog for a while, you may be familiar with a post in which I ruminated over my own responsibilities and conundrums we face in work-life balance, personal happiness, and our desires to protect ourselves or judge/shame others. And if you’ve closely followed me on Twitter or Facebook, you may have noticed we corrected a paper recently and retracted another. So I’ve stuck by my guns lately, as I long have, to correct my team’s work when I’m aware of problems. But along the way I’ve learned a lot, too, about myself, science, collaboration, humanity, how to improve research practice or scrutiny, and the pain of errors vs. the satisfaction of doing the right thing. I’ve had some excellent advice from senior management at the RVC along the way, which I am thankful for.

I’ve been realizing I should minimize my own usage of the phrase “The science may be flawed but the conclusions are right.” That can be a more-or-less valid defence, as in the case of the classic Garland paper. But it can also be a mask (unintentional or not) that hides fear that past science might have real problems (or even just minor ones that nonetheless deserve fixing) that could distract one away from the pressing issues of current science. Science doesn’t appreciate the “pay no attention to the person behind the curtain” defence, however. And we owe it to future science to tidy up past messes, ensuring the soundness of science’s data.

We’re used to moving forward in science, not backward. Indeed, the idea of moving backward, undoing one’s own efforts, can be terrifying to a scientist– especially an early career researcher, who may feel they have more at risk. But it is at the very core of science’s ethos to undo itself, to fix itself, and then to move on forward again.

I hope that this blog post inspires other scientists to think about their own research and how they balance the priorities of keeping their research chugging along but also looking backwards and reassessing it as they proceed. It should become less common to say “Yeah those data might be wrong but the conclusions are right regardless, so don’t worry.” Or it might more common to politely question such a response in others. As I wrote before, there often are no simple, one-size-fits-all answers for how to best do science. Yet that means we should be wary of letting our own simple answers slip out, lest they blind us or others.

Maybe this is all bloody obvious or tedious to blog readers but I found it interesting to think about, so I’m sharing it. I’d enjoy hearing your thoughts.

Coming soon: more Mystery Anatomy, and a Richard Owen post I’ve long intended to do.

A short post that guest-tweeting at the  Biotweeps account on Twitter got me thinking about– featuring a poll.

Imagine this: two scientists (colleagues, if you’re a scientist) are arguing thusly. Say it’s an argument about a classic paper in which much of the data subjected to detailed statistical analyses are quantitative guesses, not hard measurements. This could be in any field of science.

Scientist 1: “Conclusions are what matter most in science. If the data are guesses, but still roughly right, we shouldn’t worry much. The conclusions will still be sound regardless. That’s the high priority, because science advances by ideas gleaned from conclusions, inspiring other scientists.”

Scientist 2: “Data are what matter most in science. If the data are guesses, or flawed in some other way, this is a big problem and scientists must fix it. That’s the high priority, because science advances by data that lead to conclusions, or to more science.”

Who’s right? Have your say in this anonymous poll (please vote first before viewing results!):

link: http://poll.fm/4xf5e

[Wordpress is not showing the poll on all browsers so you may have to click the link]

And if you have more to say and don’t mind being non-anonymous, say more in the Comments- can you convince others of your answer? Or figure out what you think by ruminating in the comments?

I’m genuinely curious what people think. I have my own opinion, which has changed a lot over the past year. And I think it is a very important question scientists should think about, and discuss. I’m not just interested in scientists’ views though; anyone science-interested should join in.

[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.

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.

Even nine years later, I still keep thinking back to a day, early in my career as an academic faculty member based in England, that traumatized me. Today I’m going to share my story of that day. I feel ready to share it.

Stomach-Churning Rating: hmm that’s a tough call, but I’ll say 1/10 because it’s just photos of live crocs and such.

This day was part of a research trip that lasted a couple of weeks, and it was in Florida, not England, and little of that trip went well at first. It transpired almost exactly 9 years ago today; around 20 August 2005. I took two 2nd/3rd year undergraduate students and our lab technician with me to Florida, meeting up with Dr. Kent Vliet, an experienced crocodile specialist, to study the biomechanics of crocodile locomotion, a subject I’ve been slowwwwwwly working on since my PhD days (see recent related blog post here). We were funded by an internal grant from my university that was supposed to be seed money to get data to lay groundwork for a future large UK research grant.

Cuban crocodile adult relaxing in a nearby enclosure. Pound-for-pound, a scary croc, but these acted like puppies with their trainers.

Cuban crocodile adult relaxing in a nearby enclosure. Pound-for-pound, a scary croc, but these acted like puppies with their trainers.

I’m interested in why only some crocodylian species, of some sizes and age classes, will do certain kinds of gaits, especially mammal-like gaits such as bounding and galloping. This strongly hints at some kind of size-related biomechanical mechanism that dissuades or prevents larger crocs from getting all jiggy with it. And at large size, with few potential predators to worry about and a largely aquatic ambush predator’s ecology, why would they need to? Crocodiles should undergo major biomechanical changes in tune with their ecological shifts as they grow up. I want to know how the anatomy of crocodiles relates to these changes, and what mechanism underlies their reduction of athletic abilities like bounding. That’s the scientific motivation for working with animals that can detach limbs from your body. (The crocodiles we worked with initially on this trip were small (about 1 meter long) and not very dangerous, but they still would have done some damage if they’d chosen to bite us, and I’ve worked with a few really nasty crocs before.)

Me putting motion capture markers onto an uncooperative young Siamese crocodile.

Me putting motion capture markers onto an uncooperative young Siamese crocodile.

We worked at Gatorland (near Orlando) with some wonderfully trained crocodiles that would even sit in your lap or under your chair, and listened to vocal commands. The cuteness didn’t wear off, but our patience soon did. First, the force platform we’d borrowed (from mentor Rodger Kram’s lab; a ~$10,000 piece of useful gear) and its digital data acquisition system wouldn’t work to let us collect our data. That was very frustrating and even a very helpful local LabView software representative couldn’t solve all our problems. But at least we were able to start trying to collect data after four painstaking days of debugging while curious crocodiles and busy animal handlers waited around for us to get our act together. The stress level of our group was already mounting, and we had limited time plus plenty of real-life bugs (the bitey, itchy kind; including fire ants) and relentless heat to motivate us to get the research done.

Adorable baby Cuban crocodile.

Adorable baby Cuban crocodile.

Then the wonderfully trained crocodiles, as crocodiles will sometimes do, decided that they did not feel like doing more than a slow belly crawl over our force platform, at best. This was not a big surprise and so we patiently tried coaxing them for a couple of sweltering August days. We were working in their caged paddock, which contained a sloping grassy area, a small wooden roofed area, and then at the bottom of the slope was the crocodiles’ pond, where they sat and chilled out when they weren’t being called upon to strut their stuff for science. We didn’t get anything very useful from them, and then the weather forecast started looking ugly.

Hybrid Siamese crocodile in its pond in our enclosure, waiting to be studied.

Hybrid Siamese crocodile in its pond in our enclosure, waiting to be studied.

We’d been watching reports of a tropical storm developing off the southeastern coast of Florida, and crossing our fingers that it would miss us. But it didn’t.

When the storm hit, we were hoping to weather the edge of the storm while we packed up, because we decided we’d done our best but our time had run out and we should move to our next site, the Alligator Farm and Zoological Park in St Augustine, where I’d worked a lot before with other Crocodylia. But the storm caught us off guard, too soon, and too violently.

To give some context to the situation, for the previous several days the local croc handlers had told us stories of how lightning routinely struck this area during storms, and was particularly prone to hitting the fences on the park perimeter, which we were close to. There was a blasted old tree nearby that vultures hung out in, and they related how that blasting had been done by lightning. One trainer had been hit twice by (luckily glancing) blows from lightning hitting the fences and such.

Ominous onlooker.

Ominous onlooker.

The storm came with pounding rain and a lot of lightning, much of it clearly striking nearby- with almost no delay between flashes and thunder, and visible sky-to-ground bolts. We debated taking our forceplate out of the ground near the crocodile pond, because sensitive electrical equipment and rain don’t go well together, but this would take precious time. The forceplate was covered with a tarp to keep the rain off. I decided that, in the interest of safety, we needed to all seek shelter and let the forceplate be.

I’ll never forget the memory of leaving that crocodile enclosure and seeing a terrible sight. The crocodile pond had swiftly flooded and engulfed our forceplate. This flooding also released all the (small) crocodiles which were now happily wandering their enclosure where we’d been sitting and working before.

Another subject awaits science.

Another subject awaits science.

At that point I figured there was no going back. Lightning + deepening floodwater + electrical equipment + crocodiles = not good, so I wagered my team’s safety against our loaned equipment’s, favouring the former.

We sprinted for cars and keepers’ huts, and got split up in the rain and commotion. As the rain calmed down, I ventured out to find the rest of the team. It turned out that amidst the havoc, our intrepid lab technician had marshalled people to go fetch the forceplate out from the flooded paddock, storm notwithstanding. We quickly set to drying it out, and during some tense time over the next day we did several rounds of testing its electronics to see if it would still work. Nope, it was dead. And we still had over a week of time left to do research, but without our most useful device. (A forceplate tells you how hard animals are pushing against the ground, and with other data such as those from our motion analysis cameras, how their limbs and joints function to support them)

We went on to St Augustine and got some decent data using just our cameras, for a wide variety of crocodiles, so the trip wasn’t a total loss. I got trapped by remnants of the storm while in Washington, DC and had to sleep on chairs in Dulles Airport overnight, but I got home, totally wrecked and frazzled from the experience.

That poorly-timed storm was part of a series of powerful storms that would produce Hurricane Katrina several days later, after we’d all left Florida. So we had it relatively easy.

I’m still shaken by the experience- as a tall person who grew up in an area with a lot of dangerous storms, I was already uneasy about lightning, feeling like I had a target on my back. But running from the lightning in that storm, after all the warnings we’d had about its bad history in this area, and how shockingly close the lightning was, leaves me almost phobic about lightning strikes. I’m in awe of lightning and enjoy thunderstorms, which I’ve seen few of since I left Wisconsin in 1995, but I now hate getting caught out in them.

The ill-fated forceplate and experimental area.

The ill-fated forceplate and experimental area.

Moreover, the damage to the forceplate- which we managed to pay to repair and return to my colleague, and the failure of the Gatorland experiments, truly mortified me. I felt horrible and still feel ashamed. I don’t think I could have handled the situation much differently. It was just a shitty situation. That, and I wanted to show our undergrads a good time with research, yet what they ended up seeing was a debacle. I still have the emails I sent back to my research dean to describe what happened in the event, and they bring back the pain and stress now that I re-read them. But then… there’s a special stupid part to this story.

I tried to lighten the mood one night shortly after the storm by taking the team out to dinner, having a few drinks and then getting up to sing karaoke in front of the restaurant. I sang one of my favourite J Geil’s Band tunes- I have a nostalgic weakness for them- the song “Centerfold“. I not only didn’t sing it well (my heart was not in it and my body was shattered), and tried lamely to get the crowd involved (I think no one clapped or sang along), but also in retrospect it was a bad choice of song to be singing with two female undergrads there– I hadn’t thought about the song’s meanings when I chose to sing it, I just enjoyed it as a fun, goofy song that brought me back to innocent days of my youth in the early 1980’s. But it is not an innocent song.

So ironically, today what I feel the most embarrassed about, thinking about that whole trip and the failed experiment, is that karaoke performance. It was incredibly graceless and ill-timed and I don’t think anyone enjoyed it. I needed to unwind; the stress was crushing me; but oh… it was so damn awkward. I think I wanted to show to the team “I’m OK, I can still sing joyfully and have a good time even though we had a disastrous experiment and maybe nearly got electrified or bitten by submerged crocodiles or what-not, so you can relax too; we can move on and enjoy the rest of the trip” but in reality I proved to myself, at least, that I was not OK. And I’m still not OK about that experience. It still makes me cringe. Haunted, it took me many years to feel comfortable singing karaoke again.

It should have been a fun trip. I love working with crocodiles, but Florida is a treacherous place for field work (and many other things). I can’t say I grew stronger from this experience. There is no silver lining. It sucked, and I continually revisit it in my memory trying to find a lesson beyond “choose better times and better songs to sing karaoke with” or “stay away from floods, electricity and deadly beasts.”

So that wins, out of several good options, as the worst day(s) of my career that I can recall. I’ve had worse days in my life, but for uncomfortable science escapades this edges out some other contenders. Whenever I leave the lab to do research, I think of this experience and hope that I don’t see anything worse. It could have been much worse field work.

(Epilogue: the grants we’ve tried to fund for this crocodile gait project all got shot down, so it has lingered and we’ve done research on it gradually since, when we find time and students… And one of the students on this trip went on to do well in research and is finishing a PhD in the Structure & Motion Lab now, so we didn’t entirely scare them off science!)

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