Transitional Vertebrate Fossils FAQ
Part 1A

Contents

Part 1B

PART 1

1. Introduction

What is a transitional fossil?

The term "transitional fossil" is used at least two different ways on talk.origins, often leading to muddled and stalemated arguments. I call these two meanings the "general lineage" and the "species-to-species transition":

"General lineage":

This is a sequence of similar genera or families, linking an older group to a very different younger group. Each step in the sequence consists of some fossils that represent a certain genus or family, and the whole sequence often covers a span of tens of millions of years. A lineage like this shows obvious morphological intermediates for every major structural change, and the fossils occur roughly (but often not exactly) in the expected order. Usually there are still gaps between each of the groups -- few or none of the speciation events are preserved. Sometimes the individual specimens are not thought to be directly ancestral to the next-youngest fossils (i.e., they may be "cousins" or "uncles" rather than "parents"). However, they are assumed to be closely related to the actual ancestor, since they have intermediate morphology compared to the next-oldest and next-youngest "links". The major point of these general lineages is that animals with intermediate morphology existed at the appropriate times, and thus that the transitions from the proposed ancestors are fully plausible. General lineages are known for almost all modern groups of vertebrates, and make up the bulk of this FAQ.

"Species-to-species transition":

This is a set of numerous individual fossils that show a change between one species and another. It's a very fine-grained sequence documenting the actual speciation event, usually covering less than a million years. These species-to-species transitions are unmistakable when they are found. Throughout successive strata you see the population averages of teeth, feet, vertebrae, etc., changing from what is typical of the first species to what is typical of the next species. Sometimes, these sequences occur only in a limited geographic area (the place where the speciation actually occurred), with analyses from any other area showing an apparently "sudden" change. Other times, though, the transition can be seen over a very wide geological area. Many "species-to-species transitions" are known, mostly for marine invertebrates and recent mammals (both those groups tend to have good fossil records), though they are not as abundant as the general lineages (see below for why this is so). Part 2 lists numerous species-to-species transitions from the mammals.

Transitions to New Higher Taxa

As you'll see throughout this FAQ, both types of transitions often result in a new "higher taxon" (a new genus, family, order, etc.) from a species belonging to a different, older taxon. There is nothing magical about this. The first members of the new group are not bizarre, chimeric animals; they are simply a new, slightly different species, barely different from the parent species. Eventually they give rise to a more different species, which in turn gives rise to a still more different species, and so on, until the descendents are radically different from the original parent stock. For example, the Order Perissodactyla (horses, etc.) and the Order Cetacea (whales) can both be traced back to early Eocene animals that looked only marginally different from each other, and didn't look at all like horses or whales. (They looked rather like small, dumb foxes with raccoon-like feet and simple teeth.) But over the following tens of millions of years, the descendents of those animals became more and more different, and now we call them two different orders.

There are now several known cases of species-to-species transitions that resulted in the first members of new higher taxa. See part 2 for details.

Why do gaps exist? (or seem to exist)

Ideally, of course, we would like to know each lineage right down to the species level, and have detailed species-to-species transitions linking every species in the lineage. But in practice, we get an uneven mix of the two, with only a few species-to-species transitions, and occasionally long time breaks in the lineage. Many laypeople even have the (incorrect) impression that the situation is even worse, and that there are no known transitions at all. Why are there still gaps? And why do many people think that there are even more gaps than there really are?

Stratigraphic gaps

The first and most major reason for gaps is "stratigraphic discontinuities", meaning that fossil-bearing strata are not at all continuous. There are often large time breaks from one stratum to the next, and there are even some times for which no fossil strata have been found. For instance, the Aalenian (mid-Jurassic) has shown no known tetrapod fossils anywhere in the world, and other stratigraphic stages in the Carboniferous, Jurassic, and Cretaceous have produced only a few mangled tetrapods. Most other strata have produced at least one fossil from between 50% and 100% of the vertebrate families that we know had already arisen by then (Benton, 1989) -- so the vertebrate record at the family level is only about 75% complete, and much less complete at the genus or species level. (One study estimated that we may have fossils from as little as 3% of the species that existed in the Eocene!) This, obviously, is the major reason for a break in a general lineage. To further complicate the picture, certain types of animals tend not to get fossilized -- terrestrial animals, small animals, fragile animals, and forest-dwellers are worst. And finally, fossils from very early times just don't survive the passage of eons very well, what with all the folding, crushing, and melting that goes on. Due to these facts of life and death, there will always be some major breaks in the fossil record.

Species-to-species transitions are even harder to document. To demonstrate anything about how a species arose, whether it arose gradually or suddenly, you need exceptionally complete strata, with many dead animals buried under constant, rapid sedimentation. This is rare for terrestrial animals. Even the famous Clark's Fork (Wyoming) site, known for its fine Eocene mammal transitions, only has about one fossil per lineage about every 27,000 years. Luckily, this is enough to record most episodes of evolutionary change (provided that they occurred at Clark's Fork Basin and not somewhere else), though it misses the most rapid evolutionary bursts. In general, in order to document transitions between species, you specimens separated by only tens of thousands of years (e.g. every 20,000-80,000 years). If you have only one specimen for hundreds of thousands of years (e.g. every 500,000 years), you can usually determine the order of species, but not the transitions between species. If you have a specimen every million years, you can get the order of genera, but not which species were involved. And so on. These are rough estimates (from Gingerich, 1976, 1980) but should give an idea of the completeness required.

Note that fossils separated by more than about a hundred thousand years cannot show anything about how a species arose. Think about it: there could have been a smooth transition, or the species could have appeared suddenly, but either way, if there aren't enough fossils, we can't tell which way it happened.

Discovery of the fossils

The second reason for gaps is that most fossils undoubtedly have not been found. Only two continents, Europe and North America, have been adequately surveyed for fossil-bearing strata. As the other continents are slowly surveyed, many formerly mysterious gaps are being filled (e.g., the long-missing rodent/lagomorph ancestors were recently found in Asia). Of course, even in known strata, the fossils may not be uncovered unless a roadcut or quarry is built (this is how we got most of our North American Devonian fish fossils), and may not be collected unless some truly dedicated researcher spends a long, nasty chunk of time out in the sun, and an even longer time in the lab sorting and analyzing the fossils. Here's one description of the work involved in finding early mammal fossils: "To be a successful sorter demands a rare combination of attributes: acute observation allied with the anatomical knowledge to recognise the mammalian teeth, even if they are broken or abraded, has to be combined with the enthusiasm and intellectual drive to keep at the boring and soul-destroying task of examining tens of thousands of unwanted fish teeth to eventually pick out the rare mammalian tooth. On an average one mammalian tooth is found per 200 kg of bone-bed." (Kermack, 1984.)

Documenting a species-to-species transition is particularly grueling, as it requires collection and analysis of hundreds of specimens. Typically we must wait for some paleontologist to take it on the job of studying a certain taxon in a certain site in detail. Almost nobody did this sort of work before the mid-1970's, and even now only a small subset of researchers do it. For example, Phillip Gingerich was one of the first scientists to study species-species transitions, and it took him ten years to produce the first detailed studies of just two lineages (see part 2, primates and condylarths). In a (later) 1980 paper he said: "the detailed species level evolutionary patterns discussed here represent only six genera in an early Wasatchian fauna containing approximately 50 or more mammalian genera, most of which remain to be analyzed." [emphasis mine]

Getting the word out

There's a third, unexpected reason that transitions seem so little known. It's that even when they are found, they're not popularized. The only times a transitional fossil is noticed much is if it connects two noticably different groups (such as the "walking whale" fossil reported in 1993), or if illustrates something about the tempo and mode of evolution (such as Gingerich's work). Most transitional fossils are only mentioned in the primary literature, often buried in incredibly dense and tedious "skull & bones" papers utterly inaccessible to the general public. Later references to those papers usually collapse the known species-to-species sequences to the genus or family level. The two major college-level textbooks of vertebrate paleontology (Carroll 1988, and Colbert & Morales 1991) often don't even describe anything below the family level! And finally, many of the species-to-species transitions were described too recently to have made it into the books yet.

Why don't paleontologists bother to popularize the detailed lineages and species-to-species transitions? Because it is thought to be unnecessary detail. For instance, it takes an entire book to describe the horse fossils even partially (e.g. MacFadden's "Fossil Horses"), so most authors just collapse the horse sequence to a series of genera. Paleontologists clearly consider the occurrence of evolution to be a settled question, so obvious as to be beyond rational dispute, so, they think, why waste valuable textbook space on such tedious detail?

Misunderstanding of quotes about punctuated equilibrium

What paleontologists do get excited about are topics like the average rate of evolution. When exceptionally complete fossil sites are studied, usually a mix of patterns are seen: some species still seem to appear suddenly, while others clearly appear gradually. Once they arise, some species stay mostly the same, while others continue to change gradually. Paleontologists usually attribute these differences to a mix of slow evolution and rapid evolution (or "punctuated equilibrium": sudden bursts of evolution followed by stasis), in combination with the immigration of new species from the as-yet-undiscovered places where they first arose.

There's been a heated debate about which of these modes of evolution is most common, and this debate has been largely misquoted by laypeople, particularly creationists. Virtually all of the quotes of paleontologists saying things like "the gaps in the fossil record are real" are taken out of context from this ongoing debate about punctuated equilibrium. Actually, no paleontologist that I know of doubts that evolution has occurred, and most agree that at least sometimes it occurs gradually. The fossil evidence that contributed to that consensus is summarized in the rest of this FAQ. What they're arguing about is how often it occurs gradually. You can make up your own mind about that. (As a starting point, check out Gingerich, 1980, who found 24 gradual speciations and 14 sudden appearances in early Eocene mammals; MacFadden, 1985, who found 5 cases of gradual anagenesis, 5 cases of probable cladogenesis, and 6 sudden appearances in fossil horses; and the numerous papers in Chaline, 1983. Most studies that I've read find between 1/4-2/3 of the speciations occurring fairly gradually.)

Predictions of creationism and of evolution

Before launching into the transitional fossils, I'd like to run through the two of the major models of life's origins, biblical creationism and modern evolutionary theory, and see what they predict about the fossil record.

• Most forms of creationism hold that all "kinds" were created separately, as described in Genesis. Unfortunately there is no biological definition of "kind"; it appears to be a vague term referring to our psychological perception of types of organisms such as "dog", "tree", or "ant". In previous centuries, creationists equated "kind" to species. With the discovery of more and more evidence for derivation of one species from another, creationists bumped "kind" further up to mean higher taxonomic levels, such as "genus", or "family", though this lumps a large variety of animals in the same "kind". Some creationists say that "kind" cannot be defined in biological terms.

  Predictions of creationism: Creationists usually don't state the predictions of creationism, but I'll take a stab at it here. First, though there are several different sorts of creationism, all of them agree that there should be no transitional fossils at all between "kinds". For example, if "kind" means "species", creationism apparently predicts that there should be no species-to-species transitions whatsoever in the fossil record. If "kind" means "genus" or "family" or "order", there should be no species-to-species transitions that cross genus, family, or order lines. Furthermore, creationism apparently predicts that since life did not originate by descent from a common ancestor, fossils should not appear in a temporal progression, and it should not be possible to link modern taxa to much older, very different taxa through a "general lineage" of similar and progressively older fossils.

  Other predictions vary with the model of creationism. For instance, an older model of creationism states that fossils were created during six metaphorical "days" that may each have taken millenia to pass. This form of creationism predicts that fossils should be found in the same order outlined in Genesis: seed-bearing trees first, then all aquatic animals and flying animals, then all terrestrial animals, then humans.

  In contrast, many modern U.S. creationists believe the "Flood Theory" of the origin of fossils. The "Flood Theory" is derived from a strictly literal reading of the Bible, and states that all geological strata, and the fossils imbedded in them, were formed during the forty-day flood of Noah's time. Predictions of the Flood Theory apparently include the following:

  • most rock should be sedimentary and indicative of cataclysmic flooding. There should be no rock formations that indicate the passing of millenia of gradual accumulation of undisturbed sediment, such as multi-layered riverbed formations. There should be no large lava flows layered on top of each other, and definitely not with successively older radiometric dates in the lower levels.
  • terrestrial animal fossils should either not be sorted at all, or should be sorted by some "hydrodynamic" aspect such as body size, with, for instance, extinct elephants and large dinosaurs in the lowest layers, and small primitive dinosaurs in the upper layers. Terrestrial animal fossils should not be sorted by subtle anatomical details (such as, say, the number of cusps on the fourth premolar).
  • marine animals are a puzzle, since it is unclear that a Flood would cause any extinctions of aquatic animals. If such extinctions did occur, aquatic fossils would perhaps be "sorted" by body size or ecological niche (bottom-feeder vs. surface swimmer). For instance, plesiosaurs, primitive whales, and placoderm fishes (relatively slow-swimming and quite large) should end up in the same layers. Ichthyosaurs and porpoises (smaller, faster swimmers with almost identical body shapes and similar diets) should also occur in the same layers.
  • there should be no sorting of large rooted structures such as coral reefs and trees. There should likewise not be differential sorting of microscopic structures of the same size and shape, such as pollen grains.
  • sorting, if it occurs at all, should be quite imperfect. With only 40 days for sorting, there should be occasional examples of individual fossils that ended up in the "wrong" layer -- the occasional mammal and human fossil in Paleozoic rocks, for instance, and the occasional trilobite and plesiosaur in Cenozoic rocks.
  • sorting should not correlate with date of the surrounding rocks. If all fossils were created by Noah's flood, there is no conceivable reason that, for instance, lower layers of fossils should always end up sandwiched between lava rocks with old radiometric dates.

  Finally, some creationists believe that fossils were created by miraculous processes not operating today. (Many of these creationists combine this idea with the Flood Theory, as follows: fossils were created during the Flood, but were "sorted" by a miraculous process not observable or understandable today.) Obviously, such a theory makes no testable predictions...except perhaps for the prediction that geological formations should not bear any obvious resemblance to processes occurring today.

• Modern evolutionary theory holds that the living vertebrates arose from a common ancestor that lived hundreds of millions of years ago (via "descent with modification"; variety is introduced by mutation, genetic drift, and recombination, and is acted on by natural selection). Various proposed mechanisms of evolution differ in the expected rate and tempo of evolutionary change.

  Predictions of evolutionary theory: Evolutionary theory predicts that fossils should appear in a temporal progression, in a nested hierarchy of lineages, and that it should be possible to link modern animals to older, very different animals. In addition, the "punctuated equilibrium" model also predicts that new species should often appear "suddenly" (within 500,000 years or less) and then experience long periods of stasis. Where the record is exceptionally good, we should find a few local, rapid transitions between species. The "phyletic gradualism" model predicts that most species should change gradually throughout time, and that where the record is good, there should be many slow, smooth species-to-species transitions. These two models are not mutually exclusive -- in fact they are often viewed as two extremes of a continuum -- and both agree that at least some species-to-species transitions should be found.

What's in this FAQ

This FAQ mostly consists of a partial list of known transitions from the vertebrate fossil record. The transitions in part 1 are mostly general lineages, while in part 2 there are both general lineages and species- to-species transitions. In a hopeless attempt to save space, I concentrated almost exclusively on groups that left living descendants, ignoring all the hundreds of other groups and side-branches that have died out. I also skipped entire groups of vertebrates (most notably the dinosaurs and modern fish) in order to emphasize mammals, the group talk.origins'ers are most interested in. Note that the general lineages sometimes include "cousin" fossils. These are fossils that are thought to be very similar and closely related to the actual ancestor, but for various reasons are suspected not to be that ancestor. I have labelled them clearly in the text. I've also pointed out some of the significant remaining gaps in the vertebrate fossil record.

I got most of the information from Colbert & Morales' Evolution of the Vertebrates (1991), Carroll's Vertebrate Paleontology and Evolution (1988), Benton's The Phylogeny and Classification of the Tetrapods (1988), and from various recent papers from the scientific literature. These sources are all listed in the reference section at the end of part 2.

The time of first known appearance of each fossil is given in parentheses after the fossil name, including absolute dates when I could find them. The only exceptions are a few cases where my source didn't mention a date and it wasn't listed in Carroll's text. All of these fossils were dated by *independent* means, typically by using several different methods of radiometric dating on the strata around the fossil, and/or by cross-correlating to dated strata at other sites (e.g. MacFadden et al., 1991). The information in this FAQ assumes that these dating methods are accurate. If you have questions about the many dating methods used by paleontologists, see the other FAQs on those topics and get yourself a good textbook of sedimentary geology. Paleontologists are generally sharp cookies, and are quite persnickety about using good dating techniques.

Some terminology

"Anagenesis", "phyletic evolution":

Evolution in which an older species, as a whole, changes into a new descendent species, such that the ancestor is transformed into the descendant.

"Cladogenesis":

Evolution in which a daughter species splits off from a population of the older species, after which both the old and the young species coexist together. Notice that this allows a descendant to coexist with its ancestor.

"Chronocline":

Gradual change in one lineage over time

Ma:

Millions of years ago (a date)

my:

Millions of years (a duration)

Timescale

Summary of the known vertebrate fossil record

(We start off with primitive jawless fish.)

Transition from primitive jawless fish to sharks, skates, and rays

• Late Silurian -- first little simple shark-like denticles.
• Early Devonian -- first recognizable shark teeth, clearly derived from scales.

GAP: Note that these first, very very old traces of shark-like animals are so fragmentary that we can't get much detailed information. So, we don't know which jawless fish was the actual ancestor of early sharks.

• Cladoselache (late Devonian) -- Magnificent early shark fossils, found in Cleveland roadcuts during the construction of the U.S. interstate highways. Probably not directly ancestral to sharks, but gives a remarkable picture of general early shark anatomy, down to the muscle fibers!
• Tristychius & similar hybodonts (early Mississippian) -- Primitive proto-sharks with broad-based but otherwise shark-like fins.
• Ctenacanthus & similar ctenacanthids (late Devonian) -- Primitive, slow sharks with broad-based shark-like fins & fin spines. Probably ancestral to all modern sharks, skates, and rays. Fragmentary fin spines (Triassic) -- from more advanced sharks.
• Paleospinax (early Jurassic) -- More advanced features such as detached upper jaw, but retains primitive ctenacanthid features such as two dorsal spines, primitive teeth, etc.
• Spathobatis (late Jurassic) -- First proto-ray.
• Protospinax (late Jurassic) -- A very early shark/skate. After this, first heterodonts, hexanchids, & nurse sharks appear (late Jurassic). Other shark groups date from the Cretaceous or Eocene. First true skates known from Upper Cretaceous.

A separate lineage leads from the ctenacanthids through Echinochimaera (late Mississippian) and Similihari (late Pennsylvanian) to the modern ratfish.

Transition from from primitive jawless fish to bony fish

• Upper Silurian -- first little scales found.

GAP: Once again, the first traces are so fragmentary that the actual ancestor can't be identified.

• Acanthodians(?) (Silurian) -- A puzzling group of spiny fish with similarities to early bony fish.
• Palaeoniscoids (e.g. Cheirolepis, Mimia; early Devonian) -- Primitive bony ray-finned fishes that gave rise to the vast majority of living fish. Heavy acanthodian-type scales, acanthodian-like skull, and big notochord.
• Canobius, Aeduella (Carboniferous) -- Later paleoniscoids with smaller, more advanced jaws.
• Parasemionotus (early Triassic) -- "Holostean" fish with modified cheeks but still many primitive features. Almost exactly intermediate between the late paleoniscoids & first teleosts. Note: most of these fish lived in seasonal rivers and had lungs. Repeat: lungs first evolved in fish.
• Oreochima & similar pholidophorids (late Triassic) -- The most primitive teleosts, with lighter scales (almost cycloid), partially ossified vertebrae, more advanced cheeks & jaws.
• Leptolepis & similar leptolepids (Jurassic) -- More advanced with fully ossified vertebrae & cycloid scales. The Jurassic leptolepids radiated into the modern teleosts (the massive, successful group of fishes that are almost totally dominant today). Lung transformed into swim bladder.

Eels & sardines date from the late Jurassic, salmonids from the Paleocene & Eocene, carp from the Cretaceous, and the great group of spiny teleosts from the Eocene. The first members of many of these families are known and are in the leptolepid family (note the inherent classification problem!).

Transition from primitive bony fish to amphibians

Few people realize that the fish-amphibian transition was not a transition from water to land. It was a transition from fins to feet that took place in the water. The very first amphibians seem to have developed legs and feet to scud around on the bottom in the water, as some modern fish do, not to walk on land (see Edwards, 1989). This aquatic-feet stage meant the fins didn't have to change very quickly, the weight-bearing limb musculature didn't have to be very well developed, and the axial musculature didn't have to change at all. Recently found fragmented fossils from the middle Upper Devonian, and new discoveries of late Upper Devonian feet (see below), support this idea of an "aquatic feet" stage. Eventually, of course, amphibians did move onto the land. This involved attaching the pelvis more firmly to the spine, and separating the shoulder from the skull. Lungs were not a problem, since lungs are an ancient fish trait and were present already.

• Paleoniscoids again (e.g. Cheirolepis) -- These ancient bony fish probably gave rise both to modern ray-finned fish (mentioned above), and also to the lobe-finned fish.
• Osteolepis (mid-Devonian) -- One of the earliest crossopterygian lobe-finned fishes, still sharing some characters with the lungfish (the other lobe-finned fishes). Had paired fins with a leg-like arrangement of major limb bones, capable of flexing at the "elbow", and had an early-amphibian-like skull and teeth.
• Eusthenopteron, Sterropterygion (mid-late Devonian) -- Early rhipidistian lobe-finned fish roughly intermediate between early crossopterygian fish and the earliest amphibians. Eusthenopteron is best known, from an unusually complete fossil first found in 1881. Skull very amphibian-like. Strong amphibian- like backbone. Fins very like early amphibian feet in the overall layout of the major bones, muscle attachments, and bone processes, with tetrapod-like tetrahedral humerus, and tetrapod-like elbow and knee joints. But there are no perceptible "toes", just a set of identical fin rays. Body & skull proportions rather fishlike.
• Panderichthys, Elpistostege (mid-late Devonian, about 370 Ma) -- These "panderichthyids" are very tetrapod-like lobe-finned fish. Unlike Eusthenopteron, these fish actually look like tetrapods in overall proportions (flattened bodies, dorsally placed orbits, frontal bones! in the skull, straight tails, etc.) and have remarkably foot-like fins.
• Fragmented limbs and teeth from the middle Late Devonian (about 370 Ma), possibly belonging to Obruchevichthys -- Discovered in 1991 in Scotland, these are the earliest known tetrapod remains. The humerus is mostly tetrapod-like but retains some fish features. The discoverer, Ahlberg (1991), said: "It [the humerus] is more tetrapod-like than any fish humerus, but lacks the characteristic early tetrapod 'L-shape'...this seems to be a primitive, fish-like character....although the tibia clearly belongs to a leg, the humerus differs enough from the early tetrapod pattern to make it uncertain whether the appendage carried digits or a fin. At first sight the combination of two such extremities in the same animal seems highly unlikely on functional grounds. If, however, tetrapod limbs evolved for aquatic rather than terrestrial locomotion, as recently suggested, such a morphology might be perfectly workable."

GAP: Ideally, of course, we want an entire skeleton from the middle Late Devonian, not just limb fragments. Nobody's found one yet.

• Hynerpeton, Acanthostega, and Ichthyostega (late Devonian) -- A little later, the fin-to-foot transition was almost complete, and we have a set of early tetrapod fossils that clearly did have feet. The most complete are Ichthyostega, Acanthostega gunnari, and the newly described Hynerpeton bassetti (Daeschler et al., 1994). (There are also other genera known from more fragmentary fossils.) Hynerpeton is the earliest of these three genera (365 Ma), but is more advanced in some ways; the other two genera retained more fish- like characters longer than the Hynerpeton lineage did.
• Labyrinthodonts (eg Pholidogaster, Pteroplax) (late Dev./early Miss.) -- These larger amphibians still have some icthyostegid fish features, such as skull bone patterns, labyrinthine tooth dentine, presence & pattern of large palatal tusks, the fish skull hinge, pieces of gill structure between cheek & shoulder, and the vertebral structure. But they have lost several other fish features: the fin rays in the tail are gone, the vertebrae are stronger and interlocking, the nasal passage for air intake is well defined, etc.

More info on those first known Late Devonian amphibians: Acanthostega gunnari was very fish-like, and recently Coates & Clack (1991) found that it still had internal gills! They said: "Acanthostega seems to have retained fish-like internal gills and an open opercular chamber for use in aquatic respiration, implying that the earliest tetrapods were not fully terrestrial....Retention of fish-like internal gills by a Devonian tetrapod blurs the traditional distinction between tetrapods and fishes...this adds further support to the suggestion that unique tetrapod characters such as limbs with digits evolved first for use in water rather than for walking on land." Acanthostega also had a remarkably fish-like shoulder and forelimb. Ichthyostega was also very fishlike, retaining a fish-like finned tail, permanent lateral line system, and notochord. Neither of these two animals could have survived long on land.

Coates & Clack (1990) also recently found the first really well- preserved feet, from Acanthostega (front foot found) and Ichthyostega (hind foot found). (Hynerpeton's feet are unknown.) The feet were much more fin-like than anyone expected. It had been assumed that they had five toes on each foot, as do all modern tetrapods. This was a puzzle since the fins of lobe-finned fishes don't seem to be built on a five-toed plan. It turns out that Acanthostega's front foot had eight toes, and Ichthyostega's hind foot had seven toes, giving both feet the look of a short, stout flipper with many "toe rays" similar to fin rays. All you have to do to a lobe- fin to make it into a many-toed foot like this is curl it, wrapping the fin rays forward around the end of the limb. In fact, this is exactly how feet develop in larval amphibians, from a curled limb bud. (Also see Gould's essay on this subject, "Eight Little Piggies".) Said the discoverers (Coates & Clack, 1990): "The morphology of the limbs of Acanthostega and Ichthyostega suggest an aquatic mode of life, compatible with a recent assessment of the fish-tetrapod transition. The dorsoventrally compressed lower leg bones of Ichthyostega strongly resemble those of a cetacean [whale] pectoral flipper. A peculiar, poorly ossified mass lies anteriorly adjacent to the digits, and appears to be reinforcement for the leading edge of this paddle-like limb." Coates & Clack also found that Acanthostega's front foot couldn't bend forward at the elbow, and thus couldn't be brought into a weight-bearing position. In other words this "foot" still functioned as a horizontal fin. Ichthyostega's hind foot may have functioned this way too, though its front feet could take weight. Functionally, these two animals were not fully amphibian; they lived in an in-between fish/amphibian niche, with their feet still partly functioning as fins. Though they are probably not ancestral to later tetrapods, Acanthostega & Ichthyostega certainly show that the transition from fish to amphibian is feasible!

Hynerpeton, in contrast, probably did not have internal gills and already had a well-developed shoulder girdle; it could elevate and retract its forelimb strongly, and it had strong muscles that attached the shoulder to the rest of the body (Daeschler et al., 1994). Hynerpeton's discoverers think that since it had the strongest limbs earliest on, it may be the actual ancestor of all subsequent terrestrial tetrapods, while Acanthostega and Ichthyostega may have been a side branch that stayed happily in a mostly-aquatic niche.

In summary, the very first amphibians (presently known only from fragments) were probably almost totally aquatic, had both lungs and internal gills throughout life, and scudded around underwater with flipper-like, many-toed feet that didn't carry much weight. Different lineages of amphibians began to bend either the hind feet or front feet forward so that the feet carried weight. One line (Hynerpeton) bore weight on all four feet, developed strong limb girdles and muscles, and quickly became more terrestrial.

Transitions among amphibians

• Temnospondyls, e.g Pholidogaster (Mississippian, about 330 Ma) -- A group of large labrinthodont amphibians, transitional between the early amphibians (the ichthyostegids, described above) and later amphibians such as rhachitomes and anthracosaurs. Probably also gave rise to modern amphibians (the Lissamphibia) via this chain of six temnospondyl genera , showing progressive modification of the palate, dentition, ear, and pectoral girdle, with steady reduction in body size (Milner, in Benton 1988). Notice, though, that the times are out of order, though they are all from the Pennsylvanian and early Permian. Either some of the "Permian" genera arose earlier, in the Pennsylvanian (quite likely), and/or some of these genera are "cousins", not direct ancestors (also quite likely).
• Dendrerpeton acadianum (early Penn.) -- 4-toed hand, ribs straight, etc.
• Archegosaurus decheni (early Permian) -- Intertemporals lost, etc.
• Eryops megacephalus (late Penn.) -- Occipital condyle splitting in 2, etc.
• Trematops spp. (late Permian) -- Eardrum like modern amphibians, etc.
• Amphibamus lyelli (mid-Penn.) -- Double occipital condyles, ribs very small, etc.
• Doleserpeton annectens or perhaps Schoenfelderpeton (both early Permian) -- First pedicellate teeth! (a classic trait of modern amphibians) etc.

From there we jump to the Mesozoic:

• Triadobatrachus (early Triassic) -- a proto-frog, with a longer trunk and much less specialized hipbone, and a tail still present (but very short).
• Vieraella (early Jurassic) -- first known true frog.
• Karaurus (early Jurassic) -- first known salamander.

Finally, here's a recently found fossil:

• Unnamed proto-anthracosaur -- described by Bolt et al., 1988. This animal combines primitive features of palaeostegalians (e.g. temnospondyl-like vertebrae) with new anthracosaur-like features. Anthracosaurs were the group of large amphibians that are thought to have led, eventually, to the reptiles. Found in a new Lower Carboniferous site in Iowa, from about 320 Ma.

Contents

Part 1B