In Owen and Darwin's day, the gulf between fins and limbs seemed impossibly wide. Fish fins don't have any obvious similarities to limbs. On the outside, most fish fins are largely made up of fin webbing. Our limbs have nothing like this, nor do the limbs of any other creature alive today. The comparisons do not get any easier when you remove the fin webbing to see the skeleton inside. In most fish, there is nothing that can be compared to Owen's one bone-two bones-lotsa blobs-digits pattern. All limbs have a single long bone at their base: the humerus in the upper arm and the femur in the upper leg. In fish, the whole skeleton looks utterly different. The base of a typical fin has four or more bones inside.
In the mid-1800s, anatomists began to learn of mysterious living fish from the southern continents. One of the first was discovered by German anatomists working in South America. It looked like a normal fish, with fins and scales, but behind its throat were large vascular sacs: lungs. Yet the creature had scales and fins. So confused were the discoverers that they named the creature Lepidosiren paradoxa, "paradoxically scaled amphibian." Other fish with lungs, aptly named lungfish, were soon found in Africa and Australia. African explorers brought one to Owen. Scientists such as Thomas Huxley and the anatomist Carl Gegenbaur found lungfish to be essentially a cross between an amphibian and a fish. Locals found them delicious.
A seemingly trivial pattern in the fins of these fish had a profound impact on science. The fins of lungfish have at their base a single bone that attaches to the shoulder. To anatomists, the comparison was obvious. Our upper arm has a single bone, and that single bone, the humerus, attaches to our shoulder. In the lungfish, we have a fish with a humerus. And, curiously, it is not just any fish; it is a fish that also has lungs. Coincidence?
As a handful of these living species became known in the 1800s, clues started to come from another source. As you might guess, these insights came from ancient fish.
One of the first of these fossils came from the shores of the Gaspe Peninsula in Quebec, in rocks about 380 million years old. The fish was given a tongue-twister name, Eusthenopteron. Eusthenopteron had a surprising mix of features seen in amphibians and fish. Of Owen's one bone-two bones-lotsa blobs-digits plan of limbs, Eusthenopteron had the one bone-two bones part, but in a fin. Some fish, then, had structures like those in a limb. Owen's archetype was not a divine and eternal part of all life. It had a history, and that history was to be found in Devonian age rocks, rocks that are between 390 million and 360 million years old. This profound insight defined a whole new research program with a whole new research agenda: somewhere in the Devonian rocks we should find the origin of fingers and toes.
In the 1920s, the rocks provided more surprises. A young Swedish paleontologist, Gunnar Save-Soderbergh, was given the extraordinary opportunity to explore the east coast of Greenland for fossils. The region was terra incognita, but Save-Soderbergh recognized that it featured enormous deposits of Devonian rocks. He was one of the exceptional field paleontologists of all time, who throughout his short career uncovered remarkable fossils with both a bold exploring spirit and a precise attention to detail. (Unfortunately, he was to die tragically of tuberculosis at a young age, soon after the stunning success of his field expeditions.) In expeditions between 1929 and 1934, Save-Soderbergh's team discovered what, at the time, was labeled a major missing link. Newspapers around the world trumpeted his discovery; editorials analyzed its importance; cartoons lampooned it. The fossils in question were true mosaics: they had fish-like heads and tails, yet they also had fully formed limbs (with fingers and toes), and vertebrae that were extraordinarily amphibian-like. After Save-Soderbergh died, the fossils were described by his colleague Erik Jarvik, who named one of the new species Ichthyostega soderberghi in honor of his friend.
The fins of most fish—for example, a zebrafish (top)—have large amounts of fin webbing and many bones at the base. Lungfish captured people's interest because like us they have a single bone at the base of the appendage. Eusthenopteron (middle) showed how fossils begin to fill the gap; it has bones that compare to our upper arm and forearm. Acanthostega (bottom) shares Eusthenopteron's pattern of arm bones with the addition of fully formed digits.
For our story, Ichthyostega is a bit of a letdown. True, it is a remarkable intermediate in most aspects of its head and back, but it says very little about the origin of limbs because, like any amphibian, it already has fingers and toes. Another creature, one that received little notice when Save-Soderbergh announced it, was to provide real insights decades later. This second limbed animal was to remain an enigma until 1988, when a paleontological colleague of mine, Jenny Clack, who we introduced in the first chapter, returned to Save-Soderbergh's sites and found more of its fossils. The creature, called Acanthostega gunnari back in the 1920s on the basis of Save-Soderbergh's fragments, now revealed full limbs, with fingers and toes. But it also carried a real surprise: Jenny found that the limb was shaped like a flipper, almost like that of a seal. This suggested to her that the earliest limbs arose to help animals swim, not walk. That insight was a significant advance, but a problem remained: Acanthostega had fully formed digits, with a real wrist and no fin webbing. Acanthostega had a limb, albeit a very primitive one. The search for the origins of hands and feet, wrists and ankles had to go still deeper in time. This is where matters stood until 1995.
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