The Little Fish That Could

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I have often been teased for my habit of carrying a science book wherever I go. ("That's such a Brian thing," an acquaintance once remarked.) If I am going to be waiting for someone or have a few minutes to spare here or there I like to have something to read to fill up the time. It's either that or fiddle around with Tetris on my cell phone. Some people have told me that this manifestation of bibliophilia makes me seem antisocial,* but I cannot break the habit. I have been toting around science stuff wherever I go from a very young age.

*[My favorite instance was when I was told to "Make myself comfortable" at someone's home and was later berated for choosing to read while everyone else was milling about before lunch.]

It will come as no surprise that when I was very young I loved all things prehistoric, and one of my favorite things was a set of "Prehistoric Monsters" cards. Each card had a photograph of a model of some prehistoric animal and provided some vital statistics about the creature. I am not sure where they are now (probably in a box with all the other trading cards I amassed), but I recall them fondly, and one of the cards featured a very curious fish. Called Eusthenopteron, it was depicted as a fish that had hauled itself out of a drying pond, much like a mudskipper. (See illustration to the left.)

For most of my childhood Eusthenopteron was the icon of the evolutionary transition between fish and the earliest terrestrial vertebrates. This poor little fish, its watery home evaporated by the scorching heat, had to walk across the searing sand in the hope of finding another patch of dampness in the burning wasteland. It was this quest that gave rise to our earliest land-dwelling ancestors; if it were not for the bravery of Eusthenopteron we would not have evolved.

Today we know differently. Most of the defining features of terrestrial vertebrates, like arms, legs, wrists, fingers, &c. evolved in the water first. Creatures like Tiktaalik, Panderichthys, and Acanthostega (among many others) have helped us to better understand this transition. Eusthenopteron is still included in the transition, but as a fully-aquatic fish that represents the type of vertebrate that gave rise to "fishapods" like Tiktaalik and in turn the earliest tetrapods (four-legged vertebrates).

Indeed, scientists have learned quite a bit about the evolution of the earliest tetrapods during my lifetime, and while our present understanding is much more complete I still have a soft spot for depictions of poor little Eusthenopteron pulling itself across the drying mudflats. Where had this imagery of Eusthenopteron as an evolutionary underdog come from?

The idea that the first tetrapods evolved from fish trying to find ponds in a drying world is most often associated with the great 20th century paleontologist A.S. Romer. It was Romer's work that helped popularize this "Drying Pond Hypothesis" from the 1950's onward, but as historian Peter Bowler has demonstrated this idea had much deeper roots. Richard Swan Lull's 1917 textbook Organic Evolution provides an earlier example of the same idea.

At the time Lull was writing there were several competing hypotheses for why fish had become adapted to life on land. Perhaps fish evolved into the first tetrapods to escape predators, exploit new food resources on land, take in greater amounts of oxygen from the air, or survive the climate changes that were causing their shallow, freshwater homes to dry up. Lull favored the last of these ideas, likening the conditions the ancestors of early tetrapods faced to the dry seasons of Africa in which fish trapped in isolated polls must either go into a kind of hibernation (like lungfish) or perish.

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Lull appealed to geology to make his case. The late Devonian rock in which fish like Eusthenopteron had been found, which was considered to have preceded the evolution of the earliest tetrapods, was often the color of dried blood. This was taken as a clue by some geologists who interpreted the late Devonian strata as representing a dry, highly-seasonal world. If it was not entirely arid then the climate was sharply seasonal with a long dry period each year.

Lull hypothesized that, like living lungfish, the ancestors of tetrapods were able to preserve themselves through dry periods. They would have had lungs to survive the foul, oxygen-deprived mud that ponds dried up into, and when all moisture was gone they could go into a torpor underground until the rains returned. Eventually, however, there came a point where the dry periods were too long to survive. The fish could no longer just wait it out. If they were to survive they would have to find water on their own. Lull wrote;

The most ambitious among the lung-breathers, not content with the limitations imposed upon their lives, emerged from the age-long aquatic home and ventured into new and untried habitat. Many may have essayed the emergence, but it is probable that relentless nature, weeding out the less fit for so valorous an undertaking, destroyed all but a single sort, for there is no evidence that the ancestry of the amphibia [i.e. earliest tetrapods] is to be found in more than one evolutionary lineage.

All Lull had to work with were a probable group of bony-finned fish ancestors, a late Devonian trackway marking the appearance of early tetrapods (see above illustration), and the geological evidence. The fossils recording the transition from the water to the land were entirely missing. Those fossils, the petrified tests of Lull's hypothesis that would later be further developed by Romer, would not be found until much later.

Lull's hypothesis, and Romer's formulation of the same idea, has now been rejected in light of new evidence, yet the same imagery remains. The "fishapods" that crawled out of the shallows are often seen as heroic creatures, facing a wild world full of possibilities. If it were not for their "valorous undertaking" our species would not be here today. How this transition has fit into our expectations of "progress" within evolution, however, is a subject for another time.

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"*[My favorite instance was when I was told to "Make myself comfortable" at someone's home and was later berated for choosing to read while everyone else was milling about before lunch.]"

Me too!!!!

And you are antisocial. Embrace your antisocialness. Socialness leads to arguments, wars, death, general mayhem, etc. Social people and extroverts are the deviant ones. ;-)

By Daniel J. Andrews (not verified) on 07 Oct 2009 #permalink

^ I once got evicted from a game of Illuminati for reading little house on the prairie books. I really couldn't comment.

By stripey_cat (not verified) on 07 Oct 2009 #permalink

"The sergeant put on the poker face which has been handed down from NCO to NCO ever since one protoamphibian told another, lower ranking protoamphibian to muster a squad of newts and Take That Beach."

The best part is that walking away from a drying pond or riverbed to find another is the worst thing you can do. Staying in the deepest place is better: that's where it stays wet for the longest, and that's also what the next rain will fill up again first.

Apart from the extra-bizarre Ichthyostega, which I don't understand (and probably nobody does), it seems nowadays that not only Acanthostega but also the whatcheeriids, Crassigyrinus, colosteids, baphetids, and even most or all anthracosaurs were no more terrestrial than a catfish or eel (which sometimes come out of the water in wet nights and slither through the grass to the next body of water). The first amphibious or terrestrial vertebrates would then have appeared only shortly before the split into temnospondyls on the one hand and seymouriamorphs, lepospondyls, and diadectomorphs + amniotes on the other hand. (At least if we ignore the confusing gephyrostegids with their mutually contradictory features.)

It should also be mentioned that red beds don't require dry seasons to form -- exposure to air does seem to be necessary, but for that it's enough to be a riverbank; there is no evidence for semiarid climates in the times and places in question.

The second figure has the filename "early-tetrapod-track-thumb.jpg". Do you know where it was published, and why it was considered a tetrapod footprint? It doesn't look like one to me.

Most importantly, however, Eusthenopteron is not a "little fish". It's a meter long! :-)

By David MarjanoviÄ (not verified) on 07 Oct 2009 #permalink

Lull's emphasis on ambition and valor reminds me of Heilmann's (1925/6/7) Lamarckist idea that the origin of bird flight was basically driven by the will to fly...

By David MarjanoviÄ (not verified) on 07 Oct 2009 #permalink

David; Thanks for the comments. I wrote this mostly from a historical perspective, what Lull and others thought at the beginning of the 20th century, but I appreciate the updates all the same.

The 2nd figure is the "tetrapod track" that Lull referred to in his textbook. It should be on page 488 of Organic Evolution.

Some people have told me that this manifestation of bibliophilia makes me seem antisocial ...

Such people have an unjustified prejudice against those of us who like to read.

I seem to remember Zimmer's book, At the Water's Edge, having a similar historical perspective (complete with Eusthenopteron and Romer) in the beginning of the text and tracing the development of hypotheses as to why fish transitioned to land. One quote that stayed with me from this book was somewhere in this part where he was churning through ideas conceived and then rejected and he wrote, "Paleontology is a science of casting about."

In an interesting coincidence, I was shown a cast of that very same Devonian 'track' yesterday. David, a search for Thinopus antiquus brings up some refs.

Photos of the specimen (or casts of it) can be found here and here.

I have a book on grammar and punctuation which I read for fun when waiting for something. YANA - you are not alone.

I must tell you a joke about that one day.

I had a set of those cards too. . .what a blast from the past! I particularly remember the Brachiosaurus picture, half-submerged at the edge of a lake bed.

> Most importantly, however, Eusthenopteron is not a "little > fish". It's a meter long! :-)

And, judging by it's streamlined shape and predatory dentition, it was a fast swimmer that hunted much more formidable things than "Micro-organisms 1".

> even most or all anthracosaurs were no more terrestrial
> than a catfish or eel (which sometimes come out of the
> water in wet nights and slither through the grass to the
> next body of water). The first amphibious or terrestrial
> vertebrates would then have appeared only shortly before
> the split into temnospondyls on the one hand and
> seymouriamorphs, lepospondyls, and diadectomorphs +
> amniotes on the other hand.

Does this mean that Anthracosaurs like *Chroniosuchus* - see here: http://tanystropheus.wordpress.com/2009/06/25/wednesday-wonders-chronio… - are no longer considered to be reptilomorphs :-o?

Eusthenopteron was long pictured as the ancestral fish, just as Ichthyostega was the intrepid amphibian explorer who first ventured on to land. Paradigms have to be shattered in the light of new evidence, even when they've become cherished ideas. BTW, an old videotape series I have called "Planet Of Life" features that last image, with Ichthyostega bravely daring to crawl upon the land, the first vertebrate to do so (and curiously for a creature that had no ears, making some sort of curious loud noise!)

By Raymond Minton (not verified) on 08 Oct 2009 #permalink

My god, the nostalgia, I had those cards!

Oh god, I have that illustration.

Lets not argue and start a "newtclear war" now!

By Tim Morris (not verified) on 10 Oct 2009 #permalink

Is this card from a "snap" set or "Magic: the Gathering"? XD

By Tim Morris (not verified) on 10 Oct 2009 #permalink

I'm a little late to this thread, but ...

I'll see your Waddington's Top Trumps 'Prehistoric Monsters' from the early 1980s, and raise you with the card from my ealy childhood - the early 1970s Brooke Bond 'Prehistoric Animals' no. 1 of 50 - Eusthenopteron. Pictures of the first 9 here: http://www.brookebondcollectables.co.uk/sets/prehistoricanimals.htm and the text:

01 EUSTHENOPTERON (Greek:'strong fin')
The ancestors of the four-legged land vertebrates are to be found among the fleshy-finned fishes. The only surviving fleshy-fins, however, are the famous coelacanth (Latimeria) of the Indian Ocean and three genera of lungfish, all of which are cousins of the land vertebrates rather than their ancestors. More typical was Eusthenopteron from the Upper Devonian (350 million years ago) of North America and Europe. This carnivorous fresh-water fish, 1-2 feet long, had lobe like fins with strong muscular bases, nostrils which opened into the mouth and primitive lungs. In times of drought It could crawl on land from pool to pool in search of water and food.

http://www4.geometry.net/detail/basic_p/prehistoric_animals_dinosaurs.h…

When you consider these came free in packs of tea, marketed at the children of a family, you have to admire how the company took it seriously. They really earned their tagline "tea cards offered in the interest of education". They didn't just get some junior PR guy to run it all off in an afternoon - they employed Dr. Alan Charig of the Natural History Museum.

You wouldn't catch him calling them 'monsters' :)

Does this mean that Anthracosaurs like *Chroniosuchus* [â¦] are no longer considered to be reptilomorphs :-o?

1) Yes, the anthracosaurs are outside the temnospondyl-amniote clade according to the last few phylogenetic analyses (not all of them published yet), starting with the bootstrap tree (but not the most parsimonious trees) of Laurin & Reisz 1999 (the redescription of Solenodonsaurus).

2) It's not clear whether the chroniosuchians are anthracosaurs. The recent description of Chroniosaurus includes a phylogenetic analysis with ambiguous results.

By David MarjanoviÄ (not verified) on 07 Jan 2010 #permalink