The Fossil Record: Evolution or "Scientific Creation"

Clifford A. Cuffey

Mammal-Like Reptiles

As previously stated, a succession of transitional fossils exists that link reptiles (Class Reptilia) and mammals (Class Mammalia). These particular reptiles are classifie as Subclass Synapsida. Presently, this is the best example of th e transformation of one major higher taxon into another. The morphologic changes that took place are well documented by fossils, beginning with animals essentially 100% reptilian and resulting in animals essentially 100% mammalian. Therefore, I have chosen this as the example to summarize in more detail (Table 1, Fig. 1).

Skulls and jaws of synapsid reptiles and mammals; left column side view of skull; center column top view of skull; right column side view of lower jaw. Hylonomus modified from Carroll (1964, Figs. 2,6; 1968, Figs. 10-2, 10-5; note that Hylonomus is a protorothyrod, not a synapsid). Archaeothyris modified from Reisz (1972, Fig. 2). Haptodus modified from Currie (1977, Figs, 1a, 1b; 1979, Figs. 5a, 5b). Sphenacodo n modified from Romer & Price (1940, Fig. 4f), Allin (1975, p. 3, Fig. 16);note: Dimetrodon substituted for top view; modified from Romer & Price, 1940, pl. 10. Biarmosuchus modified from Ivakhnenko et al. (1997, pl. 65, Figs. 1a, 1B, 2); Alin & Hopson (1992; Fig. 28.4c); Sigogneau & Tchudinov (1972, Figs. 1, 15). Eoarctops modified from Broom (1932, Fig. 35a); Boonstra (1969, Fig. 18). Pristerognathus modified from Broom (1932, Figs 17a, b,c); Boonstra (1963, Fig. 5d). Procynosuchus modified from Allin & Hopson (1992, Fig. 28.4e); Hopson (1987, Fig. 5c); Brink (1963, Fig. 10a); Kemp (1979, Fig. 1); Allin (1975, p. 3, Fig. 14). Thrinaxodon modified from Allin & Hopson (1992, Fig. 28.4f);Parrington (1946, Fig. 1); Allin (1975, p. 3, Fig. 13). Probainognathus modified from Allin & Hopson (1992, Fig. 28.4g); Romer (1970, Fig. 1); Allin (1975, p. 3, Fig. 12). Morga nucodon modified from Kermack, Mussett, & Rigney (1981, Figs. 95, 99a; 1973, Fig. 7a); Allin (1975, p. 3, Fig. 11). Asioryctes modified from Carroll (1988, Fig. 20-3b). Abbreviations: ag = angular; ar = articular; cp = coronoid process; d = dentary; f = lateral temporal fenestra; j = jugal; mm = attachment site for mammalian jaw muscles; o = eye socket; po = post orbital; q = quadrate; rl = reflected lamina; sq = squamosal; ty = tympanic.

Table 1: Morphology of synapsid reptiles and mammals (Note that Hylonomus is a protothyrid, not a synapsid). Data from references cited in text.

Modern reptiles and mammals are very distinctive, easily diagnosable, and do not intergrade. Reptiles are covered by scales, mammals by hair; reptiles are cold-blooded, mammals warm-blooded; reptiles do not suckle their young, mammals have mammary glands; reptiles have sprawling posture, mammals have upright posture. Most of these features are soft part anatomy or physiology that very rarely fossilize (although dinosaur skin impressions are known from Cretaceous sediments, and imprints of mammal hair are known from Eocene bats from Germany; Franzen, 1990). In the fossil record, we must look to skeletal features.

There are many skeletal features which allow us to distinguish the reptiles from the mammals (Carroll, 1988; Table 1, rows A, M). The single most important defining characteristic is the nature of the articulation of the lower jaw to the skull (Simpson, 1959). In reptiles, multiple bones comprise the lower jaw. A small bone at the posterior end of the lower jaw, the articular, articulates with the quadrate bone of the skull (Simpson, 1959; Carroll, 1988). In mammals, one large bone, the dentary, comprises the lower jaw. It articulates with the squamosal bone of the skull (Simpson, 1959; Carroll, 1988).

From comparative anatomy studies, it is certain that most of the bones of the reptiles and mammals are homologous (Crompton & Parker, 1978; Carroll, 1988). Of greatest importance, the middle ear bones of mammals (stapes, incus, malleus, and tympanic) are homologous with several of the skull and jaw bones of reptiles (stapes, quadrate, articular, and angular, respectively; Romer, 1956, p. 33-38, 1970a; Allin, 1975, 1986; Allin & Hopson, 1992; Crompton & Parker, 1978; Hopso n, 1987, 1994; Carroll, 1988). One group of reptiles, the synapsids (Subclass Synapsida), share with the mammals an additional homologous structure: the lateral temporal fenestra, which is an opening in the skull behind the eye socket at the triple junction between the squamosal, jugal , and post orbital bones (Broom, 1932; Frazetta, 1968; Kemp, 1982; Carroll, 1988). A band of bone composed of the jugal and the squamosal is adjacent to the lateral temporal fenestra (Broom, 1932; Kemp, 1982; Carroll, 1988). This is the cheek arch so characteristic of mammal skulls (Broom, 1932; Kemp, 1982; Carroll, 1988). Therefore, synapsids are commonly named the mammal-like reptiles.

The presence of diagnosable morphologic differences between reptiles (including the oldest reptiles and the oldest synapsids) and mammals distinguishes them as distinct taxa. This allows us to test evolution by looking for transitional forms between the two. Because many of the bones are homologous, we should find evidence illustrating how these bones were modified over time to become the new bones. Furthermore, these morphologic changes should happen in parallel and in geochronologic succession.

Synapsid reptiles inhabited Pangea from the Middle Pennsylvanian through the Early Jurassic (Kemp, 1982, 1985; Sloan, 1983; Carroll, 1988; Hopson, 1969, 1987, 1994; Hopson & Crompton, 1969; Hotton, et al., 1986; Crompton & Jenkins, 1973; Sidor & Hopson, 1998; Romer & Price, 1940; Broom, 1932; Boonstra, 1963, 1969, 1971; Tchudinov, 1983; Olson, 1944; Tatarinov, 1974; Vyushkov, 1955; Efremov, 1954). From the Early Permian through the Early Triassic, they were the largest and most abundant land animals (Sloan, 1983; Colbert, 1965). Though much less well known to the general public than dinosaurs, one of the cereal box dinosaurs, Dimetrodon (the sail-backed reptile), is a synapsid, not a dinosaur (Romer & Price, 1940; Carroll, 1988). The oldest mammals are Late Triassic (Kemp, 1982; Carroll, 1988). Below is a discussion of the geochronologic succession linking synapsids and mammals. The oldest reptiles (named protorothyrids; Carroll, 1964, 1988, p. 192-199) are from the lower Middle Pennsylvanian, and the oldest synapsids (Reisz, 1972) are from the upper Middle Pennsylvanian, both of Nova Scotia. Upper Pennsylvanian and Lower Permian forms are known primarily from the midcontinent and Permian Basin region of the United States (Romer & Price, 1940; Currie, 1977, 1979; Kemp, 1982; Sloan, 1983). The basal Upper Permian forms are known from Russia (Tchudinov, 1960, 1983; Efremov, 1954; Olson, 1962; Sigogneau & Tchudinov, 1972; Ivakhnenko et al., 1997). Most of the Upper Permian and Lower Triassic succession is known from southern Africa, especially the Great Karoo of South Africa (Broom, 1932; Boonstra, 1963, 1969, 1971; Hopson & Kitching, 1972; Kemp, 1982; Sloan, 1983). The Middle Triassic forms are from South America (Romer, 1969a, 1969b, 1970b, 1973; Romer & Lewis, 1973; Bonaparte & Barbarena, 1975), and the Upper Triassic and Lower Jurassic mammals are known from Eurasia (Kermack, Mussett, & Rigney, 1973, 1981; Kemp, 1982). Subsequent Mesozoic mammals are known from all over the world (Simpson, 1928; Lillegraven et al., 1979).

When placed in proper geochronologic succession, the synapsids naturally form a succession of taxa (genera and families) that progressively become more mammal-like and less reptile-like (Kemp, 1982, 1985; Sloan, 1983; Sidor & Hopson, 1998; Hopson, 1987, 1994). Morphologic changes, summarized in Table 1 and Figure 1, affect the entire skeletal anatomy of these animals, but are most clearly displayed in their skulls.

The lateral temporal fenestra increased in size from a tiny opening smaller than the eye socket to a giant opening occupying nearly half the length of the skull. Ultimately, it merged with the eye socket, thus producing the full development of the cheek arch so characteristic of mammals (Broom, 1932; Frazetta, 1968; Kemp, 1982; Sloan, 1983; Hopson, 1987, 1994; Carroll, 1988).

Successively, the relative proportion of the lower jaw comprised of the dentary bone (teeth-bearing bone) gradually increased until the entire lower jaw consisted of the dentary (Kemp, 1982; Sloan, 1983; Carroll, 1988; Hopson, 1987, 1994). In Pennsylvanian and Lower and basal Upper Permian synapsids, the postero-dorsal edge of the lower jaw rose broadly but only slightly above the level of the tooth row (Romer & Price, 1940; Currie, 1977, 1979; Ivakhnenko et al., 1997; Tchudinov, 1960, 1983; Efremov, 1954; Olson, 1962; Sigogneau & Tchudinov, 1972; Hopson, 1987, 1994). In succeeding forms, the posterior part of the dentary expanded dorsally and posteriorly as a blade-like process, and progressively became larger (Broom, 1932; Boonstra, 1963, 1969, 1971; Sigogneau, 1970; Brink, 1963; Kemp, 1979; Hopson, 1987, 1994), forming the coronoid process (Parrington, 1946; Fourie, 1974; Romer, 1969b, 1970b, 1973; Hopson, 1987, 1994) to which the mammalian-type jaw musculature is attached (Barghusen, 1968; Bramble, 1978; Crompton, 1972; Crompton & Parker, 1978; Kemp, 1982; Sloan, 1983; Carroll, 1988). Concomitantly, the post-dentary bones progressively reduced in size (Allin, 1975; Crompton, 1972; Crompton & Parker, 1978; Kemp, 1982; Sloan, 1983; Carroll, 1988; Hopson, 1987, 1994).

Beginning with the Upper Pennsylvanian sphenacodonts, a notch developed in the angular bone that offsets a projection, the reflected lamina (Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994; Romer & Price, 1940; Currie, 1977, 1979; Kemp, 1982; Sloan, 1983; Carroll, 1988). The reflected lamina first became a large blade-like flange (Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994; Ivakhnenko et al., 1997; Tchudinov, 1960, 1983; Efremov, 1954; Olson, 1962; Sigogneau & Tchudinov, 1972; Broom, 1932; Sigogneau, 1970; Boonstra, 1963, 1969, 1971), and then was progressively reduced to a delicate horseshoe-shaped bone (Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994; Brink, 1963; Parrington, 1946; Fourie, 1974; Romer, 1969b, 1970b, 1973; Kermack, Mussett, & Rigney, 1973, 1981; Kemp, 1979, 1982; Sloan, 1983; Carroll, 1988).

Simultaneously, the quadrate progressively decreased in size (Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994; Kemp, 1982; Sloan, 1983; Carroll, 1988). The articular did not decrease in size much, being small initially, but developed a downward-pointing prong (Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994; Kemp, 1982; Sloan, 1983; Carroll, 1988). In the synapsids, the lower jaw was hinged to the skull by the articular and quadrate bones (Crompton, 1972; Crompton & Parker, 1978; Allin, 1975; Allin & Hopson, 1992; Hopson, 1987, 1994). Thus they are classified as reptiles (Simpson, 1959; Kemp, 1982; Sloan, 1983; Carroll, 1988). As the quadrate and articular became smaller, they were relieved of their solid suture to the dentary and skull (Crompton, 1972; Allin, 1975, 1986; Allin & Hopson, 1992; Hopson, 1987, 1994; Crompton & Parker, 1978; Kemp, 1982; Sloan, 1983; Carroll, 1988). A projection of the dentary extended posteriorly and made contact with the squamosal. Morganucodon possessed the mammalian dentary-squamosal jaw joint adjacent to the reptilian articular-quadrate jaw joint (Kermack, Mussett, & Rigney, 1973, 1981; Carroll, 1988). It is classified as the first mammal, but it is a perfect intermediate. Now that a new jaw joint was established, the quadrate and articular were subsequently relieved of that function (Crompton, 1972; Allin, 1975, 1986; Allin & Hopson, 1992; Hopson, 1987, 1994; Crompton & Parker, 1978; Kemp, 1982; Sloan, 1983; Carroll, 1988). Ultimately, in Middle and Upper Jurassic mammals, the tiny quadrate, articular, and ring-like angular migrated as a unit to the middle ear where they joined the stapes and became the incus, malleus, and tympanic bones (Allin, 197 5, 1986; Allin & Hopson, 1992; Hopson, 1987, 1994; Kemp, 1982; Sloan, 1983; Carroll, 1988).

Progressively, the teeth became differentiated. The large canines developed first, followed by the development of multicusped cheek teeth, reduced tooth replacement (Osborn & Crompton, 1973; Crompton & Parker, 1978), and finally full y differentiated incisors, canines, premolars, and molars with one tooth replacement during life (Kemp, 1982; Hopson, 1994).

Many other morphologic changes are documented in the fossil record. These demonstrate the morphologic and geochronologic succession from sprawling limb posture to upright limb posture of mammals (Jenkins, 1971; Romer & Lewis, 197 3; Kemp, 1982; Carroll, 1988; Hopson, 1994). As Jenkins (1971, p. 210) stated, In details of morphology and function, the cynodont post-cranial skeleton should be regarded as neither reptilian nor mammalian but as transitional between the two classes . Other changes have been adequately summarized elsewhere (Kemp, 1982; Sloan, 1983; Carroll, 1988; Hopson, 1994). Obviously, fundamental physiologic changes must have taken place as well, many of which are not directly preserved in the fossil record, though some can be inferred from the skeletal anatomy (Findlay, 1968; Kemp, 1982; Sloan, 1983, Carroll, 1988; Hopson, 1994).

This is well documented in the fossil record by a massive volume of incontrovertible data that cannot be explained away. Such large-scale, progressive, continuous, gradual, and geochronologically successive morphologic change (Sidor & Hopson, 1998) is descent with modification, and provides compelling evidence for evolution on a grand scale.