A reprint from American Scientist the magazine of Sigma Xi, The Scientific Research Society This reprint is provided for personal and noncommercial use. For any other use, please send a request to Permissions, American Scientist, P.O. Box 13975, Research Triangle Park, NC, 27709, U.S.A., or by electronic mail to [email protected]. ©Sigma Xi, The Scientific Research Society and other rightsholders Marginalia Missing Links and Found Links Pat Shipman hough missing links are of- sion enhanced by its four-to-nine-foot Tten talked about, it’s the found In and out of the length. Its skeleton differs markedly ones that hold a special place in my from those of crocodiles or alligators, heart. Found links are fossils that il- though, despite the overall resemblance lustrate major transitions during evo- water, transitional in body shape. Tiktaalik’s front fins hold lutionary history. More than that, such the biggest surprise. Each was a sort of creatures offer unexpected glimpses of forms from the fossil half-fin, half-leg containing the bony the never-predictable twists and turns elements found in a limb—with a func- taken by evolution. Their discovery record illuminate tional wrist, elbow and shoulder—and and surprise bring sheer fun to pale- yet retaining the bony “rays” of a fish ontology and biology. the nuts and bolts of fin. According to team member Farish I have always loved the iconic Arch- Jenkins, Jr., of Harvard University, the aeopteryx, a beautiful fossil recognized evolution front fins were sturdy enough to sup- in 1860 that unmistakably combines port the creature in very shallow water features of two major groups of ani- or on land for brief trips. mals: birds and reptiles. The exquisite Its broad and robust ribs were imbri- feathered wings of Archaeopteryx bear Thomas Henry Huxley as stony proof cated, like tiles on a roof. They helped most unbirdlike claws; its birdlike of evolutionary theory. Decades later, to support the body on land and prob- skull contains an avian brain but car- Archaeopteryx was trumped by an ex- ably housed lungs to supplement the ries sharp reptilian teeth, not a beak; traordinary plethora of feathered di- gills. The presence of lungs is expect- and its feathered tail is underlain by nosaurs—some nonflying—that tell ed because many of the primitive fish a long bony tail typical of a small di- different stories about the evolution of in Tiktaalik’s ancestry had lungs for nosaur, not a bird. Still, the feathers avian features. gulping air at the water’s surface as and wings on these 150-million-year- well as gills. Soft tissues are rarely old fossils qualify Archaeopteryx for the Enter the Fishapod preserved in fossils, so the lack of title of First Bird. I am equally enamored of another fossilized lungs is unremarkable. Archaeopteryx is a found link in an- found link, the fossil skeleton of Tiktaa- With or without lungs, Tiktaalik was other sense, because the anatomy of lik roseae, described on April 6, 2006, in uniquely adapted to moving between this extraordinary species reveals how the journal Nature. Tiktaalik is a name land and water. creatures evolved from propelling suggested by the elders of the Nuna- “We were absolutely surprised at the themselves along solid substrates, such vut people, who live where the fossils features of the specimens,” Ted Dae- as the ground or tree limbs, to mov- were found on Ellesmere Island in the schler of the Academy of Natural Sci- ing through the air. It was a difficult Canadian Arctic; it means “large, shal- ences, co-leader of the team, told me. transition. Archaeopteryx fascinates me low-water fish.” This 375-million-year- “That is one of the beauties of this ma- in part because its anatomy is not that old fish shows a delicious combination terial. We knew the end points—fish of a skillful, modern bird, yet it com- of unexpected features, some inherit- at the beginning and tetrapods at the peted with contemporary pterodactyls, ed from its fishy ancestors and some end—but we could not have predicted which flew using different anatomical typical of later land-dwelling tetrapods the sequence in which those anatomi- structures. I often wonder why birds (four-footed animals). Neil Shubin of cal changes occurred.” Discovering survived and those wonderful ptero- the University of Chicago, co-leader of the unexpected is one of the joys of dactyls went extinct. the discovery team, jokingly calls the paleontology. At the time of its discovery, Archae- newly discovered species a “fishapod.” A dramatic change in habitat— opteryx was hailed by the anatomist Tiktaalik’s fins, gills, scales and primi- becoming a land animal when your an- tive jaw show it was a fish. Unlike fish cestors lived in water—required many Pat Shipman is an adjunct professor of anthropol- and like tetrapods, it had a distinct anatomical changes. Sturdy limbs re- ogy at the Pennsylvania State University. Address: neck, so its head moved independently placed flexible fins. New foods had to Department of Anthropology, 315 Carpenter Build- of its body. Its flattened head and broad be found, and new means of getting ing, Pennsylvania State University, University body make Tiktaalik look somewhat them had to be developed. In this case, Park, PA 16801. Internet: [email protected] like a weird, scaly crocodile, an impres- when Tiktaalik crawled up on land it © 2006 Sigma Xi, The Scientific Research Society. Reproduction www.americanscientist.org 2006 November–December 495 with permission only. Contact [email protected]. record of fossil whales. For example, Pakicetus is a 50-million-year-old spe- cies with whalelike teeth and a whale- like skull. Its skull possesses neither the anatomical adaptations for deep diving nor those for hearing underwa- ter as well as modern whales do, sug- gesting Pakicetus used both land and shallow water environments. Most of its skeleton is still unknown, but the part of its pelvis that is known shows aquatic adaptations. Whether or not its limbs and feet were adapted for land or sea won’t be known until a more complete specimen is found. Phil Gin- gerich of the Museum of Paleontol- Ted Daeschler/VIREO ogy at the University of Michigan has Tiktaalik’s broad, flat head had eye sockets on top of the skull, making it look a lot like a found remains of Pakicetus and many crocodile, even though the creature was covered in fishy scales and had gills. To the right other spectacular whale fossils. “We of the head are the reinforced ribs and sturdy forearms that Tiktaalik used to push itself up are looking for a skeleton of Pakicetus,” from the bottom of the pond or lake. he says with a grin, emphasizing the need for an intact specimen, “knowing probably preyed upon insects. Preda- but also reveals how this specific tran- that what we find might turn out to be tory fish in the past and present often sition from water to land occurred. quite different from what we expect.” suck aquatic food into their mouths This remarkable fossil shows us Slightly younger Rodhocetus was bet- using the same mechanism that pass- something else: that the transition was ter adapted to the water, with ankles es water across the gills. But Tiktaalik not an all-or-nothing affair. “Land” or like land mammals’ that were connect- does not have a bony gill cover, which “water” is too simple a dichotomy for ed to enlarged hind feet specialized for means there was less water flow over the realities of ecosystems. There are swimming. Its front feet retained land- the gills and a less effective sucking many habitats—swamps, or shallow, adapted hooves. mechanism. Too, its snout is longer plant-choked streams, or ponds that From 45 million years ago, the fos- than in its predatory ancestors. Both shrink seasonally and occasionally dry sil whale Dorudon had a less mobile of these changes suggest that Tiktaalik up—that require a range of adaptations (more fishlike) neck, front legs modi- was snapping up prey, perhaps from to both land and water. Tiktaalik may fied into flippers, vestigial hind legs the air, rather than gulping down prey have been at home in such places. and a powerful whale tail. Dorudon along with water. shows that foot-propelled swimming Eventually tetrapods left the water Longer and Longer Swims had been superseded by the tail-pro- and relied solely on lungs for respira- Tiktaalik’s discovery made me recon- pelled swimming that characterizes tion, abandoning their gills. By simply sider the much later and opposite tran- modern whales. The features of these being, Tiktaalik not only proves that sition—from land back to the sea— three species—plus those of a dozen such major adaptive changes occurred which is documented in the excellent other related species—can be used to sketch out the way in which land spe- cies returned to the sea and reevolved their aquatic adaptations, eventually evolving into whales. Both transitions, water-to-land and land-back-to-water, occurred in a mo- saic fashion and quite possibly used those intermediate habitats to which Tiktaalik is adapted. In fish and in whales, first the shape and design of Glyptolepis Sauripterus Eusthenopteron Panderichthys Tiktaalik Acanthostega Tulerpeton the head evolved, then the forelimbs and finally the hindlimbs and tail. Are these parallels meaningful or simply coincidental? Did other lineages pre- served in the far north evolve from a fishy life to a tetrapodal one in another way? Only more analysis and more Kalliopi Monoyios fossils will tell. This diagram shows a progression of anatomical changes in the forelimb from fish to tetrapod.