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The Talk.Origins Archive: Exploring the Creation/Evolution Controversy

Transitional Vertebrate Fossils FAQ
Part 1B

Copyright © 1994-1997 by Kathleen Hunt

[Last Update: March 17, 1997]

Part 1A


Part 2A

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)

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)

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:

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)

GAP: no diapsid fossils from the mid-Permian.

Some species-to-species transitions:

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:

# Early Reptiles Mammals

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

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

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.

So, by the late Cretaceous the three groups of modern mammals were in place: monotremes, marsupials, and placentals. Placentals appear to have arisen in East Asia and spread to the Americas by the end of the Cretaceous. In the latest Cretaceous, placentals and marsupials had started to diversify a bit, and after the dinosaurs died out, in the Paleocene, this diversification accelerated. For instance, in the mid- Paleocene the placental fossils include a very primitive primate-like animal (Purgatorius - known only from a tooth, though, and may actually be an early ungulate), a herbivore-like jaw with molars that have flatter tops for better grinding (Protungulatum, probably an early ungulate), and an insectivore (Paranyctoides).

The decision as to which was the first mammal is somewhat subjective. We are placing an inflexible classification system on a gradational series. What happened was that an intermediate group evolved from the 'true' reptiles, which gradually acquired mammalian characters until a point was reached where we have artificially drawn a line between reptiles and mammals. For instance, Pachygenulus and Kayentatherium are both far more mammal-like than reptile-like, but they are both called "reptiles".

Transition from diapsid reptiles to birds

In the mid-1800's, this was one of the most significant gaps in vertebrate fossil evolution. No transitional fossils at all were known, and the two groups seemed impossibly different. Then the exciting discovery of Archeopteryx in 1861 showed clearly that the two groups were in fact related. Since then, some other reptile-bird links have been found. On the whole, though, this is still a gappy transition, consisting of a very large-scale series of "cousin" fossils. I have not included Mononychus (as it appears to be a digger, not a flier, well off the line to modern birds). See Feduccia (1980) and Rayner (1989) for more discussion of the evolution of flight, and Chris Nedin's excellent Archeopteryx FAQ for more info on that critter.

GAP: The exact reptilian ancestor of Archeopteryx, and the first development of feathers, are unknown. Early bird evolution seems to have involved little forest climbers and then little forest fliers, both of which are guaranteed to leave very bad fossil records (little animal + acidic forest soil = no remains). Archeopteryx itself is really about the best we could ask for: several specimens has superb feather impressions, it is clearly related to both reptiles and birds, and it clearly shows that the transition is feasible.

[Note: a classic study of chicken embryos showed that chicken bills can be induced to develop teeth, indicating that chickens (and perhaps other modern birds) still retain the genes for making teeth. Also note that molecular data shows that crocodiles are birds' closest living relatives.]

Part 1A


Part 2A

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