From: Subject: =?iso-8859-1?Q?Resource_of_th?= =?iso-8859-1?Q?e_American_Sci?= =?iso-8859-1?Q?entific_Affili?= =?iso-8859-1?Q?ation:_Taxonom?= =?iso-8859-1?Q?y=2C_Transitiona?= =?iso-8859-1?Q?l_Forms=2C_and_t?= =?iso-8859-1?Q?he_Fossil_Reco?= =?iso-8859-1?Q?rd_by_Keith_B.?= =?iso-8859-1?Q?_Miller?= Date: Thu, 22 Jan 2004 14:31:28 -0500 MIME-Version: 1.0 Content-Type: multipart/related; boundary="----=_NextPart_000_01C5_01C3E0F4.6C798090"; type="text/html" X-MimeOLE: Produced By Microsoft MimeOLE V5.00.3314.1001 This is a multi-part message in MIME format. ------=_NextPart_000_01C5_01C3E0F4.6C798090 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: =?iso-8859-1?B?aHR0cDovL3d3dy5hc2E=?= =?iso-8859-1?B?My5vcmcvQVNBL3Jlc28=?= =?iso-8859-1?B?dXJjZXMvTWlsbGVyLmg=?= =?iso-8859-1?B?dG1s?= Resource of the American Scientific Affiliation: = Taxonomy, Transitional Forms, and the Fossil Record by Keith B. = Miller

Taxonomy, Transitional Forms,
and the Fossil = Record


Keith B. Miller
Department of = Geology
Kansas State=20 University, Manhattan, KS 66506

The recognition and = interpretation of=20 patterns in the fossil record require an awareness of the limitations = of that=20 record. Only a very small fraction of the species that have lived = during past=20 geologic history is preserved in the rock record. Most marine species = are=20 soft-bodied, or have thin organic cuticles, and are essentially = unpreservable=20 except under the most extraordinary conditions. Furthermore, the = destructive=20 processes active in most marine environments prevent the preservation = of even=20 shelled organisms under normal conditions. Preservational = opportunities are=20 even more limited in the terrestrial environment. Most fossil = vertebrate=20 species are represented by no more than a few fragmentary remains. = Because of=20 the preservational biases of the fossil record, paleontologists must=20 reconstruct evolutionary relationships from isolated branches of an = originally=20 very bushy tree.

The process of describing and = classifying=20 organisms introduces its own patterns into the taxonomic hierarchy. = First,=20 because organisms must be placed in one group or another, taxonomy = gives the=20 impression of discontinuity. Secondly, the placement of species into = higher=20 taxa is done retrospectively; that is, by looking backward through = time. The=20 evolutionary significance of particular morphologic transitions is = only=20 recognized because of the subsequent success of particular lineages. = The=20 defining characters of higher taxa are thus a consequence of history, = and do=20 not represent some objective scale of the magnitude of morphologic = divergence.=20 Closely-related species from two different higher taxa may actually be = more=20 similar in morphology than two distantly-related species belonging to = the same=20 group.

Because new character states are = added over=20 geologic time, the morphology of species within a higher taxonomic = group=20 becomes less divergent toward the point of origin of that group. In = addition,=20 species appearing early in the history of a taxon approach more = closely the=20 morphology of species from other closely related higher taxa, often to = the=20 extent that their taxonomic assignment is uncertain. Transitional = forms=20 between higher taxa are thus a common feature of the fossil record, = although=20 continuous fossil lineages are rarely if ever preserved. Evidence from = the=20 fossil record is consistent with a wide range of proposed evolutionary = mechanisms.

Introduction

The fossil record provides persuasive evidence for macroevolutionary = change=20 and common descent. The pattern of appearance of fossil species through = geologic=20 time is critical for reconstructing evolutionary relationships. In = addition, the=20 fossil record may also contribute to our understanding of the tempo and = mode of=20 evolution, and help select between competing macroevolutionary = theories.

However, before the fossil record can be applied to these questions, = two=20 critically important topics need to be addressed. The first concerns the = completeness and resolution of the fossil record, and the second = concerns=20 taxonomic procedures. Taxonomy refers to the methods by which species = are=20 defined and grouped into a hierarchy of categories.

Nature of the Fossil Record

There are two opposite errors which need to be countered about the = fossil=20 record: (1) that it is so incomplete as to be of no value in = interpreting=20 patterns and trends in the history of life, and (2) that it is so good = that we=20 should expect a relatively complete record of the details of = evolutionary=20 transitions within most lineages.

What then is the nature of the fossil record? It can be confidently = stated=20 that only a very small fraction of the species that once lived on Earth = has been=20 preserved in the rock record and subsequently discovered and described = by=20 science. Our knowledge of the history of life can be put into = perspective by a=20 comparison with our knowledge of living organisms. About 1.5 million = living=20 species have been described by biologists, while paleontologists have = catalogued=20 only about 250,000 fossil species representing over 540 million years of = Earth=20 history (Erwin, 1993)! Why such a poor record?

Limits of the Fossil Record

Soft-bodied or thin-shelled organisms have little or no chance of=20 preservation, and the majority of species in living marine communities = are=20 soft-bodied. Consider that there are living today about 14 phyla of = worms=20 comprising nearly half of all animal phyla, yet only one, the Annelida, = has a=20 significant fossil record. The inadequacy of the fossil record to = preserve with=20 any completeness the evolutionary history of soft-bodied organisms can = be=20 illustrated by the Conodonta. Originally assigned to their own phylum, = they are=20 now believed to belong to the cordates. These soft-bodied animals are=20 represented by tiny tooth-like phosphatic fossils which are very = abundant in=20 sedimentary rocks extending over about 300 million years of Earth = history, and=20 have a worldwide distribution. Conodonts are a very important group of = marine=20 fossils for paleontologists, yet until only very recently the organism = to which=20 they belonged was completely unknown. Specimens of the worm-like = conodont animal=20 have now been discovered in Carboniferous, Ordovician, and Silurian = rocks=20 (Briggs et al., 1983; Mikulic et al., 1985; Aldridge & Purnell, = 1996). Only=20 a handful of specimens is now known from a very large and diverse group = of=20 marine animals known to be extremely abundant and widespread over a = tremendous=20 length of time!

The discovery of new soft-bodied fossil localities is always met with = great=20 enthusiasm. These localities typically turn up new species with unusual=20 morphologies, and new higher taxa are built from a few specimens! Such=20 localities are also erratically and widely spaced in geologic time = between which=20 essentially no soft-bodied fossil record exists.

Even those organisms with preservable hard parts are unlikely to be = preserved=20 under "normal" conditions. Recent studies of the fate of clam shells in = shallow=20 coastal waters reveal that shells are rapidly destroyed by scavenging, = boring,=20 chemical dissolution, and breakage. Rare events such as major storms = appear to=20 be required to incorporate shells into the sedimentary record. Getting=20 terrestrial vertebrate material into the fossil record is even more=20 difficult.

The limitations of the vertebrate fossil record can be easily = illustrated.=20 The famous fossil Archaeopteryx, occurring in a rock unit = renowned for=20 its fossil preservation, is represented by only seven known specimens, = of which=20 only two are essentially complete. Considering how many individuals of = this=20 genus probably lived and died over the thousands or millions of years of = its=20 existence, these few known specimens give some feeling for how few = individuals=20 are actually preserved as fossils and subsequently discovered. Yet this = example=20 actually represents an unusual wealth of material. The great majority of = fossil=20 vertebrate species are represented by only very fragmentary remains, and = many=20 are described on the basis of single specimens or from single = localities.=20 Complete skeletons are exceptionally rare. For many fossil taxa, = particularly=20 small mammals, the only fossils are teeth and jaw fragments. If so many = fossil=20 vertebrate species are represented by single specimens, the number of = completely=20 unknown species must be enormous!

In addition to these preservational biases, the erosion, deformation, = and=20 metamorphism of originally fossiliferous sedimentary rocks have = eliminated=20 significant portions of the fossil record over geologic time. = Furthermore, much=20 of the fossil-bearing sedimentary record is hidden in the subsurface, or = located=20 in poorly accessible or little studied geographic areas. For these = reasons, of=20 those once living species actually preserved in the fossil record, only = a small=20 portion has been discovered and described by science.

Because of the biases of the fossil record, the most abundant and=20 geographically widespread species of hard part-bearing organisms would = tend to=20 be best represented. Also, because evolutionary change is probably most = rapid=20 within small isolated populations, species within rapidly evolving = lineages are=20 less likely to be preserved in the fossil record. In addition, the = completeness=20 of the fossil record improves up the taxonomic hierarchy (Erwin, 1993). = A=20 smaller proportion of once-living species is preserved than genera, of = genera=20 than families, of families than orders, etc. As a result we can better = discern=20 the general patterns of evolutionary change than the = population-by-population or=20 species-by-species transitions.

Potential of Fossil Record for Understanding Evolutionary = Change

Given the limitations and biases discussed above, what should be = expected=20 from the fossil record? The situation is not as bleak as it may appear = from my=20 previous comments. Exceptional deposits, such as the Burgess Shale, = Solnhofen=20 Limestone, and Green River Shale, do provide surprisingly detailed = glimpses of=20 once living communities. These rare cases of exceptional preservation = (fossil=20 lagerst=E4tten) are essentially snapshots in the history of life and are = invaluable in gaining a more comprehensive picture of ancient = communities. They=20 also provide some of the most detailed anatomical data.

More commonly, thick sequences of fossiliferous rocks can enable = selected=20 skeleton or shell-bearing taxa to be examined at closely-spaced = intervals. These=20 localities provide opportunities to study patterns of evolutionary = change within=20 isolated lineages. Important information can be gained on morphologic = change=20 within species populations, and transitions between species and, rarely, = even=20 genera can be examined (Fig. 1). However, the time interval recorded by=20 continuous series of closely-spaced fossil populations is limited = because of=20 changing environmental, depositional, and preservational conditions.

Figure 1. Changes in = the shape of=20 molar teeth of the Early Eocene mammal Hypsodus, showing = evolutionary=20 transitions from species to species within a genus. (From Gingerich = [1976],=20 reprinted with permission of the American Journal of=20 Science.)

Speciation events appear to take place primarily in small isolated = peripheral=20 populations. Therefore to catch a population "in the act" requires the=20 fortuitous sampling of the particular geographic locality where the = changes=20 occurred. Even within well-preserved fossil series it is usually = difficult to=20 distinguish the record of speciation occurring within a particular = depositional=20 basin (or environment) from the effects of immigration of new species = from=20 outside that basin. For this and other reasons, well-documented and=20 widely-accepted examples of speciation in the fossil record are few (for = an=20 example, see Gingerich, 1976).

The expectation, therefore, is for the preservation of isolated = branches on=20 an originally very bushy, evolutionary tree. A few of these branches = (lineages)=20 would be fairly complete, while most are reconstructed with only very=20 fragmentary evidence (Fig. 2). While the details are missing, a general=20 understanding of the large-scale patterns and trends in evolutionary = history=20 should be discernible. Evolutionary trends over longer periods of time = and=20 across greater morphologic transitions can be followed by reconstructing = morphological sequences. Morphological transitions can be recognized in = the=20 fossil record that cross all levels of the taxonomic hierarchy.

Figure 2. The effects = of an=20 incomplete fossil record on the reconstruction of evolutionary = relationships.=20 (A) This branching tree (phylogeny) represents the actual pattern of=20 evolutionary relationships. (B) The actual preserved record of species = in the=20 fossil record might look something like this. (C) This branching tree=20 represents a possible reconstruction of the evolutionary tree based on = the=20 fossil evidence. Note that the general pattern of relationships is = preserved,=20 but that errors have been made with regard to specific = ancestor-descendant=20 relationships.

Taxonomy and Transitional Forms

Taxonomy, the process of classifying living and fossil organisms, = produces=20 its own patterns which order the diversity of life. It is thus important = to=20 recognize that names do much more than describe nature: they also = interpret it.=20 There is considerable ferment now within the field of taxonomy because = of=20 conflicting philosophies of classification, and different perceptions of = which=20 patterns in the history of life should be reflected in the taxonomic = hierarchy=20 (Eldredge & Cracraft, 1980; Schoch, 1986). Higher taxa can be either = artificial groupings of species with similar morphologies (evolutionary = grades),=20 or "natural" groups sharing derived characteristics inherited from a = common=20 ancestor (monophyletic taxa or clades).

The Linnean classification system is hierarchical, with species = grouped into=20 genera, genera into families, families into orders, etc. This system = reflects=20 the discontinuity and hierarchy observed among living organisms. = However, "this=20 system leads to the impression that species in different categories = differ from=20 one another in proportion to differences in taxonomic rank" (Carroll, = 1988, p.=20 578). This impression is false. Higher taxa are distinct and easily = recognizable=20 groups only when we ignore the time dimension of the history of life. = When the=20 fossil record is included, the boundaries between higher taxa become = blurred=20 during the major morphological radiations associated with the appearance = of new=20 higher taxa. Even in the modern world, discontinuity is not as great as = it may=20 appear superficially. In practice, species are often not easily = recognized, and=20 accepted species definitions cannot always be applied.

Another common misperception is that the origin of higher taxa does = not take=20 place at the level of populations and species. If the concept of common = descent=20 is accepted, then transitions between higher level taxonomic categories = must=20 also be species transitions (Fig. 3). This is recognized by all = evolutionary=20 paleobiologists, even those who stress the significance of the origin of = phyla=20 and classes (Valentine, 1992). Therefore, the more complete the fossil = record of=20 the origin and early radiation of higher taxa the more similar the = transitional=20 species, and the more difficult it is to determine their taxonomic = assignments.=20 Species placed into two different higher taxa may thus have very similar = morphologies.

Figure 3. Pattern of = phylogeny in=20 which one clade (or higher taxon) emerges from another. In retrospect = (time=20 T2), the two clades are seen as being distinct, and the phylogeny is = divided=20 at the position of the heavy, dashed bar into taxa A and B. A = taxonomist=20 living at time T1, however, would have recognized only a single clade = and=20 would have grouped the entire phylogeny that had developed by that = time into a=20 single taxon (A). (From Macroevolution: Pattern and Process by = Stanley=20 =A9 1979 by W.H. Freeman and Company, used with permission. All rights = reserved.)

The character states used to define higher taxa are determined=20 retrospectively. That is, they are chosen based on a knowledge of the = subsequent=20 history of the lineages possessing those traits. They do not reflect the = attainment of some objective higher level of morphologic innovation at = the time=20 of their appearance. Also, all the features subsequently identified with = a=20 particular higher taxon do not appear in a coordinated and simultaneous = manner=20 but as character mosaics within numerous closely-related species = lineages, many=20 of which are not included in the new higher taxon. In addition, as = discussed=20 above, the species associated with the origin and initial radiation of a = new=20 taxon are usually not very divergent in morphology. Were it not for the=20 subsequent evolutionary history of the lineages, species spanning the=20 transitions between families, orders, classes, and phyla would be placed = in the=20 same lower taxon (Fig. 3).

Based on the above discussion, a transitional form is simply a fossil = species=20 that possesses a morphology intermediate between that of two others = belonging to=20 different higher taxa. Such transitional forms commonly possess a = mixture of=20 traits considered characteristic of these different higher taxa. They = may also=20 possess particular characters that are themselves in an intermediate = state.=20 During the time of origin of a new higher taxon, there are often many = described=20 species with transitional morphologies representing many independent = lineages.=20 It is usually very difficult if not impossible to determine which, if = any, of=20 the known transitional forms actually lay on the lineage directly = ancestral to=20 the new taxon. For this reason, taxonomists commonly have difficulty = defining=20 higher taxa, and assigning transitional fossil species to one or the = other=20 taxon. But, although the details may elude us, the patterns of = evolutionary=20 change are in many cases well recorded in the fossil record.

Examples from the Fossil Record

As stated above, the diversity of life appears much more = discontinuous when=20 viewed at any given point in time, than it does when viewed through = time. For a=20 given time slice through the tree of life, transitions between taxa are = seen=20 only where the slice intersects the branching points of lineages. Once a = lineage=20 is split, its branches continue to evolve and diverge such that their=20 morphological (and genetic) distance increases and they become more = readily=20 distinguished taxonomic entities. When looking backward through time = using the=20 fossil record, it is found that representatives of different higher = level taxa=20 become more "primitive," that is have fewer derived characters, and = appear more=20 like the primitive members of other closely related taxa. As a result, = for=20 lineages with a good fossil record, the appearance of a new higher taxon = is=20 associated with the occurrence of species whose taxonomic identities are = uncertain or whose morphologies converge closely on that of the new = higher=20 taxon. Such patterns are found repeatedly by paleontologists.

A longstanding misperception of the fossil record of evolution is = that fossil=20 species form single lines of descent with unidirectional trends. Such a = simple=20 linear view of evolution is called orthogenesis, and has been rejected = by=20 paleontologists as a model of evolutionary change (MacFadden, 1992). The = reality=20 is much more complex than that, with numerous branching lines of descent = and=20 multiple morphologic trends (Fig. 4). The fossil record reveals that the = history=20 of life can be understood as a densely branching bush with many short = branches=20 (short-lived lineages). The well-known fossil horse series, for example, = does=20 not represent a single continuous evolving lineage (MacFadden, 1992). = Rather it=20 records more or less isolated parts of an adapting and diversifying limb = of the=20 tree of life. While incomplete, this record provides important insights = into the=20 patterns of morphological divergence and the modes of evolutionary = change.

Figure 4. Comparison = of a single=20 direct line of descent (orthogenesis) with a branching phylogeny.=20 Diversification is such an important feature of the history of life = that=20 orthogenesis is probably very rare. Fossils from a chronological = series thus=20 do not represent direct ancestor-descendant relationships, but = individual=20 branches. (From MacFadden [1992], reprinted with permission of = Cambridge=20 University Press).

Interestingly, some critics of evolution view the record of fossil = horses=20 from "Eohippus" (Hyracotherium) to Equus as trivial = (Denton, 1985). However, that is only because the intermediate forms are = known=20 (Fig. 5, 6). Without them, the morphologic distance would appear great.=20 "Eohippus" was a very small (some species only 18 inches long) = and=20 generalized herbivore (probably a browser). Besides the well-known = difference in=20 toe number (four toes at front, three at back), "Eohippus" had a = narrow=20 elongate skull with a relatively small brain and eyes forward in the = skull. It=20 possessed small canine teeth, premolars, and low-crowned simple molars. = Over=20 geologic time and within several lineages, the skull became much deeper, = the=20 eyes moved back, and the brain became larger. The incisors were widened, = premolars were altered to molars, and the molars became very = high-crowned with a=20 highly complex folding of the enamel (Evander, 1989; McFadden, = 1988).

Figure 5. Fossil = horse series=20 from Hyracotherium ("Eohippus") to Equus showing = changes=20 in skull proportions associated with an adaptive shift from browsing = to=20 grazing. This sequence shows a chronological sequence of genera within = the=20 perissodactyl family Equidae from the Eocene to the Recent. (From = MacFadden=20 [1992], reprinted with permission of Cambridge University=20 Press).

Figure 6. Stages in = horse=20 evolution showing the reduction in the number of toes and foot bones. = Forefeet=20 above, hind feet below. (A) Hyracotherium, a primitive early = Eocene=20 horse with four toes in front and three behind, (B) Miohippus, = an=20 Oligocene three-toed horse, (C) Merychippus, a late Miocene = form with=20 reduced lateral toes, and (D) Equus. (From Vertebrate=20 Paleontology by Alfred Sherwood Romer published by The University = of=20 Chicago Press, copyright =A9 1945, 1966 by The University of Chicago. = All rights=20 reserved. This material may be used and shared with the fair-use = provisions of=20 US copyright law, and it may be archived and redistributed in = electronic form,=20 provided that this entire notice, including copyright information, is = carried=20 and provided that the University of Chicago Press is notified and no = fee is=20 charged for access. Archiving, redistribution, or republication of = this text=20 on other terms, in any medium, requires both the consent of the = authors and=20 the University of Chicago Press.)

The significance of the fossil record of horses becomes clearer when = it is=20 compared with that of the other members of the order Perissodactyla = ("odd-toed=20 ungulates"). The fossil record of the extinct titanotheres is quite good = (Fig.=20 7), and the earliest representatives of this group are very similar to=20 "Eohippus" (Stanley, 1974; Mader, 1989). Likewise, the earliest = members=20 of the tapirs and rhinos were very "Eohippus"-like. Thus, the = different=20 perissodactyl groups can be traced back to a group of very similar small = generalized ungulates (Radinsky, 1979; Prothero, et al., 1989; Prothero = &=20 Schoch, 1989) (Fig. 8). But this is not all; the most primitive = ungulates=20 (hoofed mammals) are the condylarths, which are assemblages of forms=20 transitional in character between the insectivores and true ungulates = (Fig. 9).=20 Some genera and families of the condylarths had been previously assigned = to the=20 Insectivora, Carnivora, and even Primates (Romer, 1966). Thus, the = farther you=20 go back in the fossil record, the more difficult it is to place species = in their=20 "correct" higher taxonomic group. The boundaries of taxa become = blurred.

Figure 7. Stages in = the evolution=20 of the extinct perissodactyl family of the titanotheres. (A) = Eotitanops=20 (early Eocene), (B) Limnohyops (middle Eocene), (C) = Manteoceras=20 (middle Eocene), (D) Protitanotherium (late Eocene), (E)=20 Brontotherium (early Oligocene), and (F) Brontotherium. = (From=20 Stanley [1974], reprinted with permission of the journal=20 Evolution.)

Figure 8. Comparison = of the early=20 members of four perissodactyl families. (A) Hyracotherium = (Equoidea),=20 (B) Hyrachyus (Rhinoceratoidea), (C) Heptodon = ("Tapiroids"), (D)=20 Eotitanops (Titanotheriomorpha). (A and B from Vertebrate=20 Paleontology by Alfred Sherwood Romer published by The University = of=20 Chicago Press, copyright =A9 1945, 1966 by The University of Chicago. = All rights=20 reserved. This material may be used and shared with the fair-use = provisions of=20 US copyright law, and it may be archived and redistributed in = electronic form,=20 provided that this entire notice, including copyright information, is = carried=20 and provided that the University of Chicago Press is notified and no = fee is=20 charged for access. Archiving, redistribution, or republication of = this text=20 on other terms, in any medium, requires both the consent of the = authors and=20 the University of Chicago Press.) (C from Radinsky [1979], and D from = Stanley=20 [1974] both reprinted with permission of the journal=20 Evolution.)

Figure 9. (A) The = Eocene horse=20 (Hyracotherium) and representatives of the condylarths, (B)=20 Phenacodus (early Eocene) and (C) Mesonyx (middle = Eocene). Note=20 how very carnivore-like Mesonyx is although it possessed small hooves = rather=20 than claws and is classified with the ungulates. (From Vertebrate=20 Paleontology by Alfred Sherwood Romer published by The University = of=20 Chicago Press, copyright =A9 1945, 1966 by The University of Chicago. = All rights=20 reserved. This material may be used and shared with the fair-use = provisions of=20 US copyright law, and it may be archived and redistributed in = electronic form,=20 provided that this entire notice, including copyright information, is = carried=20 and provided that the University of Chicago Press is notified and no = fee is=20 charged for access. Archiving, redistribution, or republication of = this text=20 on other terms, in any medium, requires both the consent of the = authors and=20 the University of Chicago Press.)

Moving further up the taxonomic hierarchy, the condylarths and = primitive=20 carnivores (creodonts, miacids) are very similar to each other in = morphology=20 (Fig. 9, 10), and some taxa have had their assignments to these orders = changed.=20 The Miacids in turn are very similar to the earliest representatives of = the=20 Families Canidae (dogs) and Mustelidae (weasels), both of Superfamily = Arctoidea,=20 and the Family Viverridae (civets) of the Superfamily Aeluroidea. As = Romer=20 (1966) states in Vertebrate Paleontology (p. 232), "Were we = living at the=20 beginning of the Oligocene, we should probably consider all these small=20 carnivores as members of a single family." This statement also = illustrates the=20 point that the erection of a higher taxon is done in retrospect, after=20 sufficient divergence has occurred to give particular traits = significance.

Figure 10. Comparison = of skulls=20 of the early ungulates (condylarths) and carnivores. (A) The = condylarth=20 Phenacodus possessed large canines as well as cheek teeth = partially=20 adapted for herbivory. (B) The carnivore-like condylarth Mesonyx. = The=20 early Eocene creodonts (C) Oxyaena and (D) Sinopa were = primitive=20 carnivores apparently unrelated to any modern forms. (E) The Eocene=20 Vulpavus is a representative of the miacids which probably was=20 ancestral to all living carnivore groups. (From Vertebrate = Paleontology=20 by Alfred Sherwood Romer published by The University of Chicago Press, = copyright =A9 1945, 1966 by The University of Chicago. All rights = reserved. This=20 material may be used and shared with the fair-use provisions of US = copyright=20 law, and it may be archived and redistributed in electronic form, = provided=20 that this entire notice, including copyright information, is carried = and=20 provided that the University of Chicago Press is notified and no fee = is=20 charged for access. Archiving, redistribution, or republication of = this text=20 on other terms, in any medium, requires both the consent of the = authors and=20 the University of Chicago Press.)

At the level of the class, the reptile/mammal transition is = particularly well=20 documented. Near the appearance of unquestioned mammals in the fossil = record, a=20 group of mammal-like reptiles called cynodonts included species that = were=20 exceptionally mammal-like in appearance (Hopson, 1994). In skeletal = features the=20 approach to the mammalian condition was almost complete (Fig. 11, 12). = The=20 following mammalian characteristics were possessed by advanced = cynodonts: (1)=20 enlarged temporal openings with the loss of the post-orbital bar, (2) = absence of=20 the pineal eye, (3) differentiation of teeth, with front nipping teeth, = canines,=20 and molar-like back teeth, (4) a secondary palate permitting respiration = while=20 chewing, (5) a double occipital condyle which enlarges the hole for the = spinal=20 cord, (6) absence of lumbar ribs (possibly related to the presence of a=20 diaphragm), (7) a nearly erect stance, and (8) an enlarged dentary bone = in the=20 lower jaw with an extremely close approach to the mammalian jaw = articulation.=20 Furthermore, some workers argue persuasively that some mammal-like = reptiles were=20 endothermic (deRicql=E9s, 1974; Bakker, R.T., 1975; McNab, 1978). And a = few=20 exceptional fossils show evidence of glandular skin and horn (Hotton, = 1991),=20 features associated with the presence of hair.

Figure 11. = Reconstructed=20 skeletons of cynodont (advanced mammal-like reptiles) and early = mammals. (A)=20 The early Triassic cynodont Thrinaxodon and (B) the advanced = cynodont=20 Probelesodon from the middle Triassic. Note the very = mammal-like erect=20 posture of these skeletons. (C) The early mammal Megazostrodon = from the=20 early Jurassic. (All reconstructions taken from Carroll [1988], A and = C used=20 by permission of Farish A. Jenkins, Jr., Museum of Comparative = Zoology,=20 Harvard University, and B used by permission of Arnold D. Lewis, = Smithsonian=20 Institution.)

Figure 12. Comparison = of the=20 skulls of cynodonts and early mammals. The cynodont skulls are (A) the = late=20 Permian Procynosuchus; (B) the early Triassic Thrinaxodon; = (C)=20 the middle Triassic Probainognathus; and (D) the early Jurassic = Pachygenelus. Note the differentiation of the teeth and the = reduction=20 in the bones at the back of the lower jaw. The early mammal skulls are = (E) the=20 early Jurassic Sinoconodon; and (F) the early Jurassic = Morganucodon.=20 (A through D from "Systematics of the nonmammalian Synapsida and=20 implications for patterns of evolution in synapsids" by J.A. Hopson = [1991],=20 published in Origins of the Higher Groups of Tetrapods: Controversy = and=20 Consensus edited by H.-P. Schultze and L. Trueb. Used by = permission of the=20 publisher, Cornell University Press. This material is not to be = printed or=20 otherwise used without permission.) (E and F from Hopson [1994] and = used by=20 permission of James A. Hopson.)

The complex of transitional fossil forms has created significant = problems for=20 the definition of the class Mammalia (Desui, 1991). For most workers, = the=20 establishment of a squamosal-dentary jaw articulation is considered one = of the=20 primary defining characters. The transition in jaw articulation from = reptiles to=20 mammals is particularly illustrative of the appearance of a class level=20 morphologic character (Fig. 12). In reptiles, the lower jaw contains = several=20 bones, and the articular bone at the back of the jaw articulates with = the=20 quadrate bone of the skull. In mammals, the lower jaw has only one bone, = the=20 dentary, and it articulates with the squamosal bone of the skull. Within = the=20 cynodont lineage, the dentary bone becomes progressively larger and the = other=20 bones are reduced to nubs at the back. In one group of advanced = cynodonts, the=20 dentary bone has been brought nearly into contact with the squamosal, = and in=20 another, a secondary articulation exists between the surangular (another = small=20 bone at the back of the jaw) and squamosal (Hopson, 1991). The earliest = known=20 mammals, the morganucodonts, retain the vestigial lower jaw bones of the = reptiles. These small bones still form a reduced, but functional, = reptilian jaw=20 joint medial to the new dentary-squamosal mammalian articulation. These=20 reptilian jaw elements were subsequently detached completely from the = jaw to=20 become the mammalian middle ear (Crompton & Parker, 1978). Better=20 intermediate character states could hardly be imagined!

As with most transitions between higher taxonomic categories, there = is more=20 than one lineage that possesses intermediate morphologies. Again, this = is=20 consistent with both the expectations of evolutionary theory, and the = nature of=20 the fossil record. The prediction would be for a bush of many lineages, = many of=20 which would be dead ends. Because of their objective to erect only = monophyletic=20 taxa (an ancestor is grouped with all of its descendants), some = paleontologists=20 have advocated including mammals with the advanced cynodonts, or even = with the=20 whole group of mammal-like reptiles, in a single higher taxon (Desui, = 1991).

As in the case of the reptile-mammal transition, the distinctiveness = of the=20 classes also becomes blurred during the amphibian-reptile transition. = The oldest=20 known reptiles (Fig. 13) have been collected within the fossilized = stumps of=20 lycopod trees from the late Pennsylvanian in Nova Scotia (Carroll, 1970, = 1991).=20 Several groups of reptiliomorph amphibians occur near the appearance of = these=20 unquestioned reptiles. Some of these (the seymouriamorphs and = diadectomorphs)=20 were in fact previously regarded as reptiles (Carroll, 1988; Benton, = 1991).

Figure 13. Skeleton = and skull of=20 the earliest known reptile Hylonomus from the early = Pennsylvanian.=20 Reptiliomorph amphibians placed in a group called the anthracosaurs = converge=20 closely on the reptiles in skeletal morphology (see reconstructions of = the=20 anthracosaur amphibians Bruktererpeton and Proterogyrinus = in=20 Carroll [1991]). (From Vertebrate Paleontology by Alfred = Sherwood Romer=20 published by The University of Chicago Press, copyright =A9 1945, 1966 = by The=20 University of Chicago. All rights reserved. This material may be used = and=20 shared with the fair-use provisions of US copyright law, and it may be = archived and redistributed in electronic form, provided that this = entire=20 notice, including copyright information, is carried and provided that = the=20 University of Chicago Press is notified and no fee is charged for = access.=20 Archiving, redistribution, or republication of this text on other = terms, in=20 any medium, requires both the consent of the authors and the = University of=20 Chicago Press.)

Fossil Transitions Associated with Major Adaptive Shifts

Of special interest in the history of life are the morphological = transitions=20 associated with the major adaptive shifts from water to land, land to = water, and=20 land to air. These major changes in mode of life opened up tremendous = new=20 adaptive opportunities for animal life. While the fossil evidence for = some of=20 these transitions is minimal, for others exciting parts of the puzzle = have been=20 uncovered.

The transition from water to land was one of the most significant = events in=20 animal evolution. Recent paleontological and systematic work has shed = new light=20 on this transition (Fig. 14). The most primitive amphibian yet known is = the late=20 Devonian Ichthyostega, a tetrapod with a flattened skull and = bearing a=20 tail fin. The limbs were until recently poorly known, but new fossil = evidence=20 has come to light. The hand, previously unknown, shows that these = amphibians=20 possessed seven to eight digits. The limbs also had a very limited range = of=20 movement and the animal was not as well adapted to terrestrial = locomotion as=20 previously thought (Ahlberg & Milner, 1994). The rhipidistian fishes = are=20 widely considered to have given rise to the amphibians. One small group = of late=20 Devonian rhipidistians, the panderichthyids, appears to be closely = related to=20 the ichthyostegids (Schultze, 1991). These fishes have flattened skulls = very=20 similar to that of the early amphibians. In addition, the anal and = dorsal fins=20 are absent, and the tail is very similar to that of Ichthyostega=20 (Vorobyeva & Schultze, 1991). The lobed pectoral and pelvic fins = have bones=20 that homologize with the limb bones of the tetrapods. Whether part of a = single=20 direct lineage or not, ichthyostegid amphibians and panderichthyid = fishes are=20 clearly transitional forms between class level taxa. The first known = skull of a=20 panderichthyid was in fact initially considered to be an amphibian = (Vorobyeva=20 & Schultze, 1991), again illustrating the taxonomic problems = encountered=20 during the appearance and early radiation of a new taxon.

Figure 14. The transition from fish to amphibian = illustrated=20 by body form and skeletons, with details of skulls and vertebrae. (A)=20 Osteolepiform fish Eusthenopteron; (B) panderichthyid fish=20 Panderichthys; and (C) labyrinthodont amphibian = Ichthyostega.=20 (From Ahlberg & Milner [1994], reprinted with permission from=20 Nature, copyright =A9 1994 Macmillan Magazines Limited, and = from Per=20 Ahlberg.)

Probably one of the most celebrated and mysterious transitions has = been that=20 of the origin of whales from a primitive condylarth (ungulate) ancestor. = The=20 earliest whales possessed skulls similar in many ways to those of a = group of=20 Eocene carnivorous condylarths called mesonycids. Until 1993 the = earliest fossil=20 whales were only known from partial skulls with no postcranial material. = However, several very important transitional fossils from Pakistan have = been=20 described over the last several years (Gingerich, et al., 1993) and more = discoveries are certain to follow. The geologically oldest included = enough of=20 the skeleton to reveal that this otter-sized whale had short front limbs = and=20 longer hind legs with large feet apparently used in swimming (Berta, = 1994;=20 Thewissen, et al., 1994). The second, somewhat younger species had = shorter hind=20 limbs indicating a trend toward reduction in limb size (Gingerich, et = al.,=20 1994). Whales apparently evolved in what is now Pakistan since all the = known=20 fossil material for earliest whales has been found in that geographic = area.=20 Because the evolution of new body plans is likely to occur in an = isolated=20 geographic area, the discovery of the fossil record of such transitions = is=20 dependent on the serendipitous sampling of the right locality.

The most famous of transitional fossils is the earliest known bird,=20 Archeopteryx. Ostrum has described over 20 shared characteristics = between=20 Archeopteryx and coelurosaur theropods. Among these are: a = theropod-like=20 pelvis, the close similarities of the bones of the forelimbs including a = swivel=20 wrist joint, and the similarity of the hind limbs and feet with the = presence of=20 a reversed first toe (Hecht, et al., 1985; Dodson, 1985; Ostrom, 1994). = The=20 similarities of Archeopteryx to theropod dinosaurs such as=20 Velociraptor and Deinonychus are especially strong, and a = newly=20 discovered dinosaur called Unenlagia has features of the limbs = and pelvis=20 that are the most bird-like yet known (Novas & Puerta, 1997). As = interesting=20 as the similarities with the theropods are, the differences between=20 Archeopteryx and modern birds are also significant: it has a long = bony=20 tail, a sternum is absent, its vertebrae are not fused together over the = pelvis=20 to form a synsacrum, and air ducts are absent in its long bones. In most = respects, Archeopteryx is more of a flying feathered dinosaur = than a=20 bird. In the last several years the discovery of new fossil birds from = the=20 Cretaceous has led to the erection of a whole new subclass of primitive = birds=20 called the enantiornithes (Chiappe, 1995). This new group includes = several=20 fossil species previously identified as theropod dinosaurs (e.g.,=20 Ornithomimus)! There are also some newly discovered fossils whose = classification as theropod or bird is in dispute (Chiappe, 1995). The = recent=20 discovery in China of a theropod dinosaur with the possible preservation = of fine=20 feathers, even suggests that feathers may not be exclusively = characteristic of=20 birds (Morell, 1997). This again illustrates the taxonomic uncertainties = that=20 surround transitional forms.

Conclusions

From this brief survey of fossil vertebrates, it is clear that = transitional=20 forms between higher taxa are common features of the fossil record. The=20 morphology of species within a higher taxonomic group becomes less = divergent=20 toward the point of origin of that group. Morphological diversity and = disparity=20 increase with time. In addition, transitional species possess mixtures = of=20 morphologic characters from different higher taxa often to the extent = that their=20 taxonomic assignment is uncertain. This pattern is obscured by taxonomy = which=20 gives a false impression of discontinuity.

The fossil record thus provides good evidence for the large-scale = patterns=20 and trends in evolutionary history. Recognizing its limitations, the = fossil=20 record appears to be consistent with the wide range of evolutionary = mechanisms=20 already proposed. Any wholesale abandonment of present paradigms would = be very=20 premature. Many critical gaps in our knowledge remain, but as evident = from this=20 review important discoveries are continually being made that intrigue, = surprise,=20 and enrich our understanding of the evolutionary history of life.

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