Evidence for turtle evolution
by Jerry Bergman and Wayne Frair
The fossil record is rich with many well-preserved turtle shells and a wide variety
of turtles ‘dating back’ to before the dinosaurs in the Triassic. Phylogenetic
analysis of turtles has resulted in much controversy and conflicting results that
vary depending on the techniques used. Molecular evidence also contradicts previous
evolutionary classifications based on gross morphological comparisons. The extant
evidence shows that turtles appear in the fossil record fully formed.
Image by Lisa Pizzarella.
An artist’s conception of Proganochelys based on fossil evidence. Note the
distinct mountains and valleys in the top shell. The top shell and other features
are very similar to the modern alligator turtle.
Turtles are ideal animals for testing evolutionary ideas because some of their most
unique structures, such as their hard shells, preserve extremely well in the fossil
record. What also makes turtles perfect candidates for studying evolution is that
their body plan is unique among tetrapods, and would require ‘some remarkable
changes in the skeleton and internal organs’ as they evolved from a typical
tetrapod to a carapace-plastron clad turtle.1,2 An example is that the scapula
of vertebrates is outside the rib cage, but in turtles the scapula, the humerus
and several other bones are all inside of the rib cage.3 Moreover, breathing is very different compared with
other reptiles because the turtle chest is not distensible.4 Therefore, if turtle evolution were true, we would
expect the fossil record to provide a better record of intermediate morphologies
than for many other vertebrates. However, the evidence for such ‘missing links’
is still missing.
The fossil record
With Proganochelys [the ‘oldest’ known turtle], our trail into the past
runs cold. We do not know from whence it came.—evolutionary turtle expert
R. Orenstein.
Extensive fossil turtle deposits extending back to the Triassic5 have been found throughout the world, including
Germany, India, Thailand, South Africa, North America and China.6 The earliest known turtle, named Proganochelys
and discovered in Germany in the 1880s, was dated by evolutionists to 210–220
million years ago. The specimens had a shell consisting of 60 plates of various
sizes, and a carapace up to 1 m long. Its skeleton was ‘characteristic of
turtles—carapace, plastron, scapular girdle inside the rib cage (unique among
vertebrates).’7
This primarily aquatic turtle possessed ‘cervical vertebrae with well developed
acuminate, spiny apophyses—making it impossible for the turtle to retract
its head’ to protect itself.8
Most known modern turtles can retract their heads (a major exception are sea turtles).
No intermediate forms between these first turtles ‘and any other reptile,
living or extinct’ has ever been found:
‘With Proganochelys, our trail into the past runs cold. We do not
know from whence it came. We may be not be much closer to knowing today than we
were more than a century ago, in the 1880s, when Proganochelys was first
discovered … the possible choices for the original turtle span almost the
entire range of reptiles, living and extinct.’9
Proganochelys ‘literally pops into the fossil record as a completely
formed turtle.’10
Furthermore the whole chelonian body plan ‘appears in the fossil record without
intermediates, and the relationship of turtles to other amniote groups is not certain.’11 Gilbert et al.
concluded that the ‘absence of intermediates or transitional forms in the
fossil record’, especially when the fossil record is coupled with the developmental
and anatomical novelties exhibited by turtles, argues that turtles arose saltationally.12 The late well-known paleontologist
George Gaylord Simpson emphasized a well known fact that the fossil record of most
taxa, including turtles,
‘ … appear abruptly. … A great many sequences of two or a few
temporally intergrading species are known, but even at this level most species appear
without known immediate ancestors, and really long, perfectly complete
sequences of numerous species are exceedingly rare.13
The abundant turtle fossil record supports the conclusion that turtles have remained
‘unchanged for at least 150 million years.’14 Flank concludes that ‘Turtle fossils are
found more often than other animals of similar size, and the evolutionary history
of the modern turtle is fairly well known’ except the earliest turtle ancestors
and, as a result, ‘the exact ancestry of living turtles is disputed among
paleontologists.’15
Monophyletic or polyphyletic evolution
Another question evolutionists are unable to answer is: did the various families
of chelonians evolve from some common ancestor or by parallel or convergent evolution?
One theory is that sea turtles evolved from land turtles, requiring significant
evolutionary changes to adapt to the sea. For example, sea turtles filter salt from
sea water by producing large salty tears. Feet must evolve into flippers, requiring
extremely elongated phalanges. Yet not one transitional sea turtle fossil has been
found.16,17 However, the consensus among herpetologists is
that
‘ … the evolutionary position of turtles with the amniote phylogeny
has eluded evolutionary biologists for more than a century. This phylogenetic problem
has remained unsolved partly because turtles have such a unique morphology that
only few characters can be used to link them with any other group of amniotes.’18
Proganochelys ‘literally pops into the fossil record as a completely formed
turtle.’—J.R. Spotila
Another hypothesis postulates that modern turtles evolved from the Chelidae, a ‘primitive’
side necked turtle unique to Australia and South America.19 Other herpetologists argue for a placodont ancestor,
especially a Henodus because of its turtle-like appearance. Since there
are many phylogenic problems of postulating turtle evolution from a Henodus,
others speculate that the similarity of turtles and Henodus is explained
by convergent evolution.20
As Rieppel and Reisz conclude, ‘Turtle relationships remain labile, and further
investigations of their relationships are required.’20
Many evolutionary trees exist, all based largely on speculation rather than fossil
evidence. So far, no convincing evidence exists for any view.
Evolution of the turtle shell
[The] problem for an evolutionary biologist is to explain these transformations
in the context of a gradualistic process.—O. Rieppel, who supports a ‘hopeful
monster’ explanation
Because of the lack of fossil intermediates, evolutionists have to resort to speculative
hypotheses to rescue to fit turtles into evolution. One hypothesis is that the turtle
carapace gradually evolved from ‘elements of the primitive reptilian integument.’21 Reptile expert Olivier
Rieppel argued that a big ‘problem for an evolutionary biologist is to explain
these transformations in the context of a gradualistic process.’3
Rieppel argues that turtles could not evolve by a gradual process, and concluded
that they may be an example of ‘hopeful monsters’.3
More recently Gilbert and his associates22
have proposed a theoretical embryological model involving movement of the ribs into
the dermal layer leading to the evolution of a turtle shell. This modeling, although
useful, cannot replace the need for paleontological evidence.23
Turtle teeth
It is also theorized that ancient turtles possessed teeth but lost them. A good
example is that Proganochelys had midpalatal homodont ‘teeth’
which were actually small denticles formed by the development of a tough covering
over some of the bones of the palate (which modern turtles lack), yet was otherwise
similar to modern turtles. The evolution of teeth is the problem evolutionists have
to deal with and, conversely, the loss of teeth would be expected in the biblical
model of the Fall which predicts corruption of the genome and the accompanying deterioration
of the phenotype. A mutation could easily have occurred in one of the ‘tooth’
development genes in turtles that disabled tooth maturation but still allowed the
animal to survive. If it proved beneficial for its specific feeding habits, it may
even be selected for as a result. Loss of teeth has evidently occurred several times
in history in animals, including possibly some birds, and certain monotremes such
as the platypus.
Moreover, this provides no solace for evolutionists because such a process results
in a loss of genetic information, not the addition of completely new information
that the evolution of turtles from their putative ancestors requires. Because of
their genetic constitution turtles have produced a great range of morphologies.
Their wide genetic variation allows the creation of variety both through careful
breeding and by various natural mechanisms. Moreover, fossilized turtles prove that
far greater chelonian diversity existed in the past than is found today.24
Biochemical comparisons
Researchers, disappointed with the lack of progress in understanding turtle evolution,
have increasingly looked to molecular and physiological studies to solve this evolutionary
enigma.25 However, comparisons
of turtles have conflicted with the hypothesized phylogenies based on gross morphological
comparisons.26
Hedges and Poling assembled all of the known genetic data available in order to
resolve the controversy. According to their analysis of the largest available collection
of reptile genes, turtles have been found by genetic studies to be the closest relatives
of, not birds as once believed, but rather of crocodiles and alligators. They concluded
that ‘The results provide strong evidence that the turtle is the crocodile’s
closest living relative.’27
‘The study’s conclusions contradict decades of research based on anatomical
and fossil studies, which had firmly positioned birds as the reptile group most
closely related to crocodiles and alligators, a group known as crocodilians.’27
Other researchers have concluded that molecular data favours the view that archosaurs
(crocodiles and birds) are
‘ … the living sister group of turtles, whereas morphological studies
support lepidosaurs (tuatara, lizards, and snakes) as the closest living relatives
of turtles. Accepting these hypotheses implies that turtles cannot be viewed any
longer as primitive reptiles, and that they might have lost the temporal holes in
the skull secondarily rather than never having had them.’18
After a study of nuclear DNA-coded proteins, Iwabe, et al. (2005) also
concluded that turtles belong to a monophyletic cluster including birds and crocodiles.
They emphatically stated that, ‘All other possible tree topologies were significantly
rejected.’28
In summary, the molecular research has, so far, provided evidence to support the
conclusion that ‘the molecular data conflict with paleontological data …
and it will be a challenge not only to paleontologists … but also to molecular
systematists to resolve these conflicts.’29
Increasing research has tended to uncover more conflicts between molecular and gross
morphological data as time passes.
Conclusions
Turtles are an ideal life form for evaluating the evidence for evolution, not only
because their shells are readily preserved but also because of the abundance of
turtles in the fossil record. Yet no evidence ever has been found in support of
the evolution of turtles from a non-turtle ancestor in spite of over a century of
extensive fossil explorations and the identification of many thousands of fossils.
‘Turtles are so different from any other reptile that their peculiarities
are practically useless as a guide for distinguishing among potential ancestors,
and the origin of turtles remains one of the great unanswered questions of evolutionary
biology … the possible choices for the original turtle span almost the entire
range of reptiles, living and extinct.’9
The origin of turtles has long been, and continues to be, a major evolutionary enigma.
The oldest known turtles clearly were turtles. Since turtles appear abruptly in
the fossil record, the current data are consistent with a creation event followed
by considerable diversification coupled with degeneration.
Acknowledgments
We wish to thank Eric Blievernicht, Jody Allen and Clifford Lillo for their helpful
comments.
Related articles
Further reading
Related resources
References
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- Rieppel, O., Turtles as hopeful monsters, BioEssays
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- Bellairs, ref. 2, p. 66. Return to text.
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