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Transitional Fossils

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Posted by Mario Dovalina on May 16, 2002 05:01:53 UTC

For your reading pleasure, a report on transitional fossils from talkorigins.org.

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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 rapidest 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
CENOZOIC

(See part 2) 65-0 Ma Mammals & birds & teleost fish dominant

MESOZOIC

Cretaceous 144-65 Ma Dinosaurs dominant. Small mammals, birds.

Jurassic 213-144 Ma Dinosaurs dominant. First mammals, then first birds.

Triassic 248-213 Ma Mammalian reptiles dominant. First dinosaurs.

PALEOZOIC

Permian 286-248 Ma Amphibians dominant. First mammal-like reptiles.

Pennsylvanian 320-286 Ma Amphibians dominant. First reptiles.

Mississippian 360-320 Ma Big terrestrial amphibians, fishes.

Devonian 408-360 Ma Fish dominant. First amphibians.

Silurian 438-408 Ma First ray-finned & lobe-finned fish.

Ordovician 505-438 Ma More jawless fishes.

Cambrian 590-505 Ma First jawless fishes.


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.

Transition from amphibians to amniotes (first reptiles)
The major functional difference between the ancient, large amphibians and the first little reptiles is the amniotic egg. Additional differences include stronger legs and girdles, different vertebrae, and stronger jaw muscles. For more info, see Carroll (1988) and Gauthier et al. (in Benton, 1988)

Proterogyrinus or another early anthracosaur (late Mississippian) -- Classic labyrinthodont-amphibian skull and teeth, but with reptilian vertebrae, pelvis, humerus, and digits. Still has fish skull hinge. Amphibian ankle. 5-toed hand and a 2-3-4-5-3 (almost reptilian) phalangeal count.
Limnoscelis, Tseajaia (late Carboniferous) -- Amphibians apparently derived from the early anthracosaurs, but with additional reptilian features: structure of braincase, reptilian jaw muscle, expanded neural arches.
Solenodonsaurus (mid-Pennsylvanian) -- An incomplete fossil, apparently between the anthracosaurs and the cotylosaurs. Loss of palatal fangs, loss of lateral line on head, etc. Still just a single sacral vertebra, though.
Hylonomus, Paleothyris (early Pennsylvanian) -- These are protorothyrids, very early cotylosaurs (primitive reptiles). They were quite little, lizard-sized animals with amphibian-like skulls (amphibian pineal opening, dermal bone, etc.), shoulder, pelvis, & limbs, and intermediate teeth and vertebrae. Rest of skeleton reptilian, with reptilian jaw muscle, no palatal fangs, and spool-shaped vertebral centra. Probably no eardrum yet. Many of these new "reptilian" features are also seen in little amphibians (which also sometimes have direct-developing eggs laid on land), so perhaps these features just came along with the small body size of the first reptiles.
The ancestral amphibians had a rather weak skull and paired "aortas" (systemic arches). The first reptiles immediately split into two major lines which modified these traits in different ways. One line developed an aorta on the right side and strengthened the skull by swinging the quadrate bone down and forward, resulting in an enormous otic notch (and allowed the later development of good hearing without much further modification). This group further split into three major groups, easily recognizable by the number of holes or "fenestrae" in the side of the skull: the anapsids (no fenestrae), which produced the turtles; the diapsids (two fenestrae), which produced the dinosaurs and birds; and an offshoot group, the eurapsids (two fenestrae fused into one), which produced the ichthyosaurs.

The other major line of reptiles developed an aorta on left side only, and strengthened the skull by moving the quadrate bone up and back, obliterating the otic notch (making involvement of the jaw essential in the later development of good hearing). They developed a single fenestra per side. This group was the synapsid reptiles. They took a radically different path than the other reptiles, involving homeothermy, a larger brain, better hearing and more efficient teeth. One group of synapsids called the "therapsids" took these changes particularly far, and apparently produced the mammals.

Some transitions among reptiles
I will review just a couple of the reptile phylogenies, since there are so many.... Early reptiles to turtles: (Also see Gaffney & Meylan, in Benton 1988)

Captorhinus (early-mid Permain) -- Immediate descendent of the protorothryids.
Here we come to a controversy; there are two related groups of early anapsids, both descended from the captorhinids, that could have been ancestral to turtles. Reisz & Laurin (1991, 1993) believe the turtles descended from procolophonids, late Permian anapsids that had various turtle-like skull features. Others, particularly Lee (1993) think the turtle ancestors are pareiasaurs:

Scutosaurus and other pareiasaurs (mid-Permian) -- Large bulky herbivorous reptiles with turtle-like skull features. Several genera had bony plates in the skin, possibly the first signs of a turtle shell.
Deltavjatia vjatkensis (Permian) -- A recently discovered pareiasaur with numerous turtle-like skull features (e.g., a very high palate), limbs, and girdles, and lateral projections flaring out some of the vertebrae in a very shell-like way. (Lee, 1993)
Proganochelys (late Triassic) -- a primitive turtle, with a fully turtle-like skull, beak, and shell, but with some primitive traits such as rows of little palatal teeth, a still-recognizable clavicle, a simple captorhinid-type jaw musculature, a primitive captorhinid- type ear, a non-retractable neck, etc..
Recently discovered turtles from the early Jurassic, not yet described.
Mid-Jurassic turtles had already divided into the two main groups of modern turtles, the side-necked turtles and the arch-necked turtles. Obviously these two groups developed neck retraction separately, and came up with totally different solutions. In fact the first known arch-necked turtles, from the Late Jurassic, could not retract their necks, and only later did their descendents develop the archable neck. Early reptiles to diapsids: (see Evans, in Benton 1988, for more info)

Hylonomus, Paleothyris (early Penn.) -- The primitive amniotes described above
Petrolacosaurus, Araeoscelis (late Pennsylvanian) -- First known diapsids. Both temporal fenestra now present. No significant change in jaw muscles. Have Hylonomus-style teeth, with many small marginal teeth & two slightly larger canines. Still no eardrum.
Apsisaurus (early Permian) -- A more typical diapsid. Lost canines. (Laurin, 1991)
GAP: no diapsid fossils from the mid-Permian.

Claudiosaurus (late Permian) -- An early diapsid with several neodiapsid traits, but still had primitive cervical vertebrae & unossified sternum. probably close to the ancestry of all diapsides (the lizards & snakes & crocs & birds).
Planocephalosaurus(early Triassic) -- Further along the line that produced the lizards and snakes. Loss of some skull bones, teeth, toe bones.
Protorosaurus, Prolacerta (early Triassic) -- Possibly among the very first archosaurs, the line that produced dinos, crocs, and birds. May be "cousins" to the archosaurs, though.
Proterosuchus (early Triassic) -- First known archosaur.
Hyperodapedon, Trilophosaurus (late Triassic) -- Early archosaurs.
Some species-to-species transitions:

De Ricqles (in Chaline, 1983) documents several possible cases of gradual evolution (also well as some lineages that showed abrupt appearance or stasis) among the early Permian reptile genera Captorhinus, Protocaptorhinus, Eocaptorhinus, and Romeria.
Horner et al. (1992) recently found many excellent transitional dinosaur fossils from a site in Montana that was a coastal plain in the late Cretaceous. They include:
Many transitional ceratopsids between Styracosaurus and Pachyrhinosaurus
Many transitional lambeosaurids (50! specimens) between Lambeosaurus and Hypacrosaurus.
A transitional pachycephalosaurid between Stegoceras and Pachycephalosaurus
A transitional tyrannosaurid between Tyrannosaurus and Daspletosaurus.
All of these transitional animals lived during the same brief 500,000 years. Before this site was studied, these dinosaur groups were known from the much larger Judith River Formation, where the fossils showed 5 million years of evolutionary stasis, following by the apparently abrupt appearance of the new forms. It turns out that the sea level rose during that 500,000 years, temporarily burying the Judith River Formation under water, and forcing the dinosaur populations into smaller areas such as the site in Montana. While the populations were isolated in this smaller area, they underwent rapid evolution. When sea level fell again, the new forms spread out to the re-exposed Judith River landscape, thus appearing "suddenly" in the Judith River fossils, with the transitional fossils only existing in the Montana site. This is an excellent example of punctuated equilibrium (yes, 500,000 years is very brief and counts as a "punctuation"), and is a good example of why transitional fossils may only exist in a small area, with the new species appearing "suddenly" in other areas. (Horner et al., 1992) Also note the discovery of Ianthosaurus, a genus that links the two synapsid families Ophiacodontidae and Edaphosauridae. (see Carroll, 1988, p. 367)

Transition from synapsid reptiles to mammals
This is the best-documented transition between vertebrate classes. So far this series is known only as a series of genera or families; the transitions from species to species are not known. But the family sequence is quite complete. Each group is clearly related to both the group that came before, and the group that came after, and yet the sequence is so long that the fossils at the end are astoundingly different from those at the beginning. As Rowe recently said about this transition (in Szalay et al., 1993), "When sampling artifact is removed and all available character data analyzed [with computer phylogeny programs that do not assume anything about evolution], a highly corroborated, stable phylogeny remains, which is largely consistent with the temporal distributions of taxa recorded in the fossil record." Similarly, Gingerich has stated (1977) "While living mammals are well separated from other groups of animals today, the fossil record clearly shows their origin from a reptilian stock and permits one to trace the origin and radiation of mammals in considerable detail." For more details, see Kermack's superb and readable little book (1984), Kemp's more detailed but older book (1982), and read Szalay et al.'s recent collection of review articles (1993, vol. 1).

This list starts with pelycosaurs (early synapsid reptiles) and continues with therapsids and cynodonts up to the first unarguable "mammal". Most of the changes in this transition involved elaborate repackaging of an expanded brain and special sense organs, remodeling of the jaws & teeth for more efficient eating, and changes in the limbs & vertebrae related to active, legs-under-the-body locomotion. Here are some differences to keep an eye on:


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# Early Reptiles Mammals


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1 No fenestrae in skull - Massive fenestra exposes all of braincase

2 Braincase attached loosely - Braincase attached firmly to skull

3 No secondary palate - Complete bony secondary palate

4 Undifferentiated dentition - Incisors, canines, premolars, molars

5 Cheek teeth uncrowned points - Cheek teeth (PM & M) crowned & cusped

6 Teeth replaced continuously - Teeth replaced once at most

7 Teeth with single root - Molars double-rooted

8 Jaw joint quadrate-articular - Jaw joint dentary-squamosal (*)

9 Lower jaw of several bones - Lower jaw of dentary bone only

10 Single ear bone (stapes) - Three ear bones (stapes, incus, malleus)

11 Joined external nares - Separate external nares

12 Single occipital condyle - Double occipital condyle

13 Long cervical ribs - Cervical ribs tiny, fused to vertebrae

14 Lumbar region with ribs - Lumbar region rib-free

15 No diaphragm - Diaphragm

16 Limbs sprawled out from body - Limbs under body

17 Scapula simple - Scapula with big spine for muscles

18 Pelvic bones unfused - Pelvis fused

19 Two sacral (hip) vertebrae - Three or more sacral vertebrae

20 Toe bone #'s 2-3-4-5-4 - Toe bones 2-3-3-3-3

21 Body temperature variable - Body temperature constant


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(*) The presence of a dentary-squamosal jaw joint has been arbitrarily selected as the defining trait of a mammal.

Paleothyris (early Pennsylvanian) -- An early captorhinomorph reptile, with no temporal fenestrae at all.
Protoclepsydrops haplous (early Pennsylvanian) -- The earliest known synapsid reptile. Little temporal fenestra, with all surrounding bones intact. Fragmentary. Had amphibian-type vertebrae with tiny neural processes. (reptiles had only just separated from the amphibians)
Clepsydrops (early Pennsylvanian) -- The second earliest known synapsid. These early, very primitive synapsids are a primitive group of pelycosaurs collectively called "ophiacodonts".
Archaeothyris (early-mid Pennsylvanian) -- A slightly later ophiacodont. Small temporal fenestra, now with some reduced bones (supratemporal). Braincase still just loosely attached to skull. Slight hint of different tooth types. Still has some extremely primitive, amphibian/captorhinid features in the jaw, foot, and skull. Limbs, posture, etc. typically reptilian, though the ilium (major hip bone) was slightly enlarged.
Varanops (early Permian) -- Temporal fenestra further enlarged. Braincase floor shows first mammalian tendencies & first signs of stronger attachment to rest of skull (occiput more strongly attached). Lower jaw shows first changes in jaw musculature (slight coronoid eminence). Body narrower, deeper: vertebral column more strongly constructed. Ilium further enlarged, lower-limb musculature starts to change (prominent fourth trochanter on femur). This animal was more mobile and active. Too late to be a true ancestor, and must be a "cousin".
Haptodus (late Pennsylvanian) -- One of the first known sphenacodonts, showing the initiation of sphenacodont features while retaining many primitive features of the ophiacodonts. Occiput still more strongly attached to the braincase. Teeth become size-differentiated, with biggest teeth in canine region and fewer teeth overall. Stronger jaw muscles. Vertebrae parts & joints more mammalian. Neural spines on vertebrae longer. Hip strengthened by fusing to three sacral vertebrae instead of just two. Limbs very well developed.
Dimetrodon, Sphenacodon or a similar sphenacodont (late Pennsylvanian to early Permian, 270 Ma) -- More advanced pelycosaurs, clearly closely related to the first therapsids (next). Dimetrodon is almost definitely a "cousin" and not a direct ancestor, but as it is known from very complete fossils, it's a good model for sphenacodont anatomy. Medium-sized fenestra. Teeth further differentiated, with small incisors, two huge deep- rooted upper canines on each side, followed by smaller cheek teeth, all replaced continuously. Fully reptilian jaw hinge. Lower jaw bone made of multiple bones & with first signs of a bony prong later involved in the eardrum, but there was no eardrum yet, so these reptiles could only hear ground-borne vibrations (they did have a reptilian middle ear). Vertebrae had still longer neural spines (spectacularly so in Dimetrodon, which had a sail), and longer transverse spines for stronger locomotion muscles.
Biarmosuchia (late Permian) -- A therocephalian -- one of the earliest, most primitive therapsids. Several primitive, sphenacodontid features retained: jaw muscles inside the skull, platelike occiput, palatal teeth. New features: Temporal fenestra further enlarged, occupying virtually all of the cheek, with the supratemporal bone completely gone. Occipital plate slanted slightly backwards rather than forwards as in pelycosaurs, and attached still more strongly to the braincase. Upper jaw bone (maxillary) expanded to separate lacrymal from nasal bones, intermediate between early reptiles and later mammals. Still no secondary palate, but the vomer bones of the palate developed a backward extension below the palatine bones. This is the first step toward a secondary palate, and with exactly the same pattern seen in cynodonts. Canine teeth larger, dominating the dentition. Variable tooth replacement: some therocephalians (e.g Scylacosaurus) had just one canine, like mammals, and stopped replacing the canine after reaching adult size. Jaw hinge more mammalian in position and shape, jaw musculature stronger (especially the mammalian jaw muscle). The amphibian-like hinged upper jaw finally became immovable. Vertebrae still sphenacodontid-like. Radical alteration in the method of locomotion, with a much more mobile forelimb, more upright hindlimb, & more mammalian femur & pelvis. Primitive sphenacodontid humerus. The toes were approaching equal length, as in mammals, with #toe bones varying from reptilian to mammalian. The neck & tail vertebrae became distinctly different from trunk vertebrae. Probably had an eardrum in the lower jaw, by the jaw hinge.
Procynosuchus (latest Permian) -- The first known cynodont -- a famous group of very mammal-like therapsid reptiles, sometimes considered to be the first mammals. Probably arose from the therocephalians, judging from the distinctive secondary palate and numerous other skull characters. Enormous temporal fossae for very strong jaw muscles, formed by just one of the reptilian jaw muscles, which has now become the mammalian masseter. The large fossae is now bounded only by the thin zygomatic arch (cheekbone to you & me). Secondary palate now composed mainly of palatine bones (mammalian), rather than vomers and maxilla as in older forms; it's still only a partial bony palate (completed in life with soft tissue). Lower incisor teeth was reduced to four (per side), instead of the previous six (early mammals had three). Dentary now is 3/4 of lower jaw; the other bones are now a small complex near the jaw hinge. Jaw hinge still reptilian. Vertebral column starts to look mammalian: first two vertebrae modified for head movements, and lumbar vertebrae start to lose ribs, the first sign of functional division into thoracic and lumbar regions. Scapula beginning to change shape. Further enlargement of the ilium and reduction of the pubis in the hip. A diaphragm may have been present.
Dvinia [also "Permocynodon"] (latest Permian) -- Another early cynodont. First signs of teeth that are more than simple stabbing points -- cheek teeth develop a tiny cusp. The temporal fenestra increased still further. Various changes in the floor of the braincase; enlarged brain. The dentary bone was now the major bone of the lower jaw. The other jaw bones that had been present in early reptiles were reduced to a complex of smaller bones near the jaw hinge. Single occipital condyle splitting into two surfaces. The postcranial skeleton of Dvinia is virtually unknown and it is not therefore certain whether the typical features found at the next level had already evolved by this one. Metabolic rate was probably increased, at least approaching homeothermy.
Thrinaxodon (early Triassic) -- A more advanced "galesaurid" cynodont. Further development of several of the cynodont features seen already. Temporal fenestra still larger, larger jaw muscle attachments. Bony secondary palate almost complete. Functional division of teeth: incisors (four uppers and three lowers), canines, and then 7-9 cheek teeth with cusps for chewing. The cheek teeth were all alike, though (no premolars & molars), did not occlude together, were all single- rooted, and were replaced throughout life in alternate waves. Dentary still larger, with the little quadrate and articular bones were loosely attached. The stapes now touched the inner side of the quadrate. First sign of the mammalian jaw hinge, a ligamentous connection between the lower jaw and the squamosal bone of the skull. The occipital condyle is now two slightly separated surfaces, though not separated as far as the mammalian double condyles. Vertebral connections more mammalian, and lumbar ribs reduced. Scapula shows development of a new mammalian shoulder muscle. Ilium increased again, and all four legs fully upright, not sprawling. Tail short, as is necessary for agile quadrupedal locomotion. The whole locomotion was more agile. Number of toe bones is 2.3.4.4.3, intermediate between reptile number (2.3.4.5.4) and mammalian (2.3.3.3.3), and the "extra" toe bones were tiny. Nearly complete skeletons of these animals have been found curled up - a possible reaction to conserve heat, indicating possible endothermy? Adults and juveniles have been found together, possibly a sign of parental care. The specialization of the lumbar area (e.g. reduction of ribs) is indicative of the presence of a diaphragm, needed for higher O2 intake and homeothermy. NOTE on hearing: The eardrum had developed in the only place available for it -- the lower jaw, right near the jaw hinge, supported by a wide prong (reflected lamina) of the angular bone. These animals could now hear airborne sound, transmitted through the eardrum to two small lower jaw bones, the articular and the quadrate, which contacted the stapes in the skull, which contacted the cochlea. Rather a roundabout system and sensitive to low-frequency sound only, but better than no eardrum at all! Cynodonts developed quite loose quadrates and articulars that could vibrate freely for sound transmittal while still functioning as a jaw joint, strengthened by the mammalian jaw joint right next to it. All early mammals from the Lower Jurassic have this low-frequency ear and a double jaw joint. By the middle Jurassic, mammals lost the reptilian joint (though it still occurs briefly in embryos) and the two bones moved into the nearby middle ear, became smaller, and became much more sensitive to high-frequency sounds.
Cynognathus (early Triassic, 240 Ma; suspected to have existed even earlier) -- We're now at advanced cynodont level. Temporal fenestra larger. Teeth differentiating further; cheek teeth with cusps met in true occlusion for slicing up food, rate of replacement reduced, with mammalian-style tooth roots (though single roots). Dentary still larger, forming 90% of the muscle-bearing part of the lower jaw. TWO JAW JOINTS in place, mammalian and reptilian: A new bony jaw joint existed between the squamosal (skull) and the surangular bone (lower jaw), while the other jaw joint bones were reduced to a compound rod lying in a trough in the dentary, close to the middle ear. Ribs more mammalian. Scapula halfway to the mammalian condition. Limbs were held under body. There is possible evidence for fur in fossil pawprints.
Diademodon (early Triassic, 240 Ma; same strata as Cynognathus) -- Temporal fenestra larger still, for still stronger jaw muscles. True bony secondary palate formed exactly as in mammals, but didn't extend quite as far back. Turbinate bones possibly present in the nose (warm-blooded?). Dental changes continue: rate of tooth replacement had decreased, cheek teeth have better cusps & consistent wear facets (better occlusion). Lower jaw almost entirely dentary, with tiny articular at the hinge. Still a double jaw joint. Ribs shorten suddenly in lumbar region, probably improving diaphragm function & locomotion. Mammalian toe bones (2.3.3.3.3), with closely related species still showing variable numbers.
Probelesodon (mid-Triassic; South America) -- Fenestra very large, still separate from eyesocket (with postorbital bar). Secondary palate longer, but still not complete. Teeth double-rooted, as in mammals. Nares separated. Second jaw joint stronger. Lumbar ribs totally lost; thoracic ribs more mammalian, vertebral connections very mammalian. Hip & femur more mammalian.
Probainognathus (mid-Triassic, 239-235 Ma, Argentina) -- Larger brain with various skull changes: pineal foramen ("third eye") closes, fusion of some skull plates. Cheekbone slender, low down on the side of the eye socket. Postorbital bar still there. Additional cusps on cheek teeth. Still two jaw joints. Still had cervical ribs & lumbar ribs, but they were very short. Reptilian "costal plates" on thoracic ribs mostly lost. Mammalian #toe bones.
Exaeretodon (mid-late Triassic, 239Ma, South America) -- (Formerly lumped with the herbivorous gomphodont cynodonts.) Mammalian jaw prong forms, related to eardrum support. Three incisors only (mammalian). Costal plates completely lost. More mammalian hip related to having limbs under the body. Possibly the first steps toward coupling of locomotion & breathing. This is probably a "cousin" fossil not directly ancestral, as it has several new but non-mammalian teeth traits.
GAP of about 30 my in the late Triassic, from about 239-208 Ma. Only one early mammal fossil is known from this time. The next time fossils are found in any abundance, tritylodontids and trithelodontids had already appeared, leading to some very heated controversy about their relative placement in the chain to mammals. Recent discoveries seem to show trithelodontids to be more mammal- like, with tritylodontids possibly being an offshoot group (see Hopson 1991, Rowe 1988, Wible 1991, and Shubin et al. 1991). Bear in mind that both these groups were almost fully mammalian in every feature, lacking only the final changes in the jaw joint and middle ear.

Oligokyphus, Kayentatherium (early Jurassic, 208 Ma) -- These are tritylodontids, an advanced cynodont group. Face more mammalian, with changes around eyesocket and cheekbone. Full bony secondary palate. Alternate tooth replacement with double-rooted cheek teeth, but without mammalian-style tooth occlusion (which some earlier cynodonts already had). Skeleton strikingly like egg- laying mammals (monotremes). Double jaw joint. More flexible neck, with mammalian atlas & axis and double occipital condyle. Tail vertebrae simpler, like mammals. Scapula is now substantially mammalian, and the forelimb is carried directly under the body. Various changes in the pelvis bones and hind limb muscles; this animal's limb musculature and locomotion were virtually fully mammalian. Probably cousin fossils (?), with Oligokyphus being more primitive than Kayentatherium. Thought to have diverged from the trithelodontids during that gap in the late Triassic. There is disagreement about whether the tritylodontids were ancestral to mammals (presumably during the late Triassic gap) or whether they are a specialized offshoot group not directly ancestral to mammals.
Pachygenelus, Diarthrognathus (earliest Jurassic, 209 Ma) -- These are trithelodontids, a slightly different advanced cynodont group. New discoveries (Shubin et al., 1991) show that these animals are very close to the ancestry of mammals. Inflation of nasal cavity, establishment of Eustachian tubes between ear and pharynx, loss of postorbital bar. Alternate replacement of mostly single- rooted teeth. This group also began to develop double tooth roots -- in Pachygenelus the single root of the cheek teeth begins to split in two at the base. Pachygenelus also has mammalian tooth enamel, and mammalian tooth occlusion. Double jaw joint, with the second joint now a dentary-squamosal (instead of surangular), fully mammalian. Incipient dentary condyle. Reptilian jaw joint still present but functioning almost entirely in hearing; postdentary bones further reduced to tiny rod of bones in jaw near middle ear; probably could hear high frequencies now. More mammalian neck vertebrae for a flexible neck. Hip more mammalian, with a very mammalian iliac blade & femur. Highly mobile, mamma

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