Over a hundred
authors, including
molecular phylogeneticists, systematists,
and paleontologists, contributed
to The Timetree of Life,
which its editors say is the first volume
to publish calibrated divergence
times against phylogenies
for all major groups of living
things. The results are truly impressive
reviews of the histories and
current knowledge of molecular
and other determinations of when
the major groups of living things
diverged from each other.
In assembling this book, the
authors submitted to certain conventions.
Alternate phylogenies are
not considered. Only timetrees
based on molecules are used. The
editors say that only one kind of
molecule needs to be used, and
that it doesn’t have to be independently
validated by other (including
non-molecular) lines of evidence.
If different molecules give
different divergence estimates, the
estimates are to be averaged. (This
strikes me as strange, inasmuch as
it equally values or doubts all studies,
rather than asking questions
about the reliability of certain molecules
or studies over others.) In
fact, the editors in particular seem
to be glossing over a lot of legitimate
debate and cognitive dissonance,
which seems odd for a scientific
book.
For example, in their opening
chapter the editors reject fossil evidence
as reliable for estimating
divergence times. Instead, they
advocate “associated geological
dates,” an approach first proposed
by Charles Sibley in the 1970s and
quickly discredited. (What is the
“date” of the opening of the
Atlantic Ocean?) They give the
emergence of islands as good
events by which to calibrate divergence
times. But what major
groups of organisms first diverged
on islands? The editors actually
don’t give a single example of a
reliable “geologic event” that can
calibrate molecular phylogenies.
The editors state that “fossil calibrations
are always minimum
times of divergence,” which would
be true if one simply used the first
recorded appearance of a member
of a stem or node group. More reliable
is the assessment of the timing
of appearances of characters
that are diagnostic of that group.
Let’s say that node group A shares
derived features 1–5, and that its
immediate relatives can be recognized
because they have progressive
subsets of 1–5 (for example,
one critter has 1, another has 1 &
2, and so on). Knowing when these
immediate relatives lived provides
very strong control on divergence
dates based on fossils. Of course,
the fossil record may not be good
enough to decide these questions
in the great majority of cases. On
the other hand, unconstrained
extrapolations from molecular differentiation
rates, with no independent
lines of evidence to test
them, are technologically impressive
but empirically unsatisfactory.
It is interesting that the other
introductory chapters disagree
with the editors’ methods. John
Avise forthrightly extols the use of
fossils to calibrate divergence
times. Gradstein and Ogg lay out
the geologic time scale and the
important certainties and uncertainties
in its calibration. Benton,
Donoghue, and Asher, all paleontologists,
in a particularly impressive
review (with over 500 references)
show how using both fossils and
molecules in tandem can produce
reliable results for much of the
phylogeny of Metazoa.
So the editors seem to be broadminded
in including eclectic
approaches to assembling the
timetrees of life. It is too bad, then,
that the prescriptions of Benton
and others are not followed
throughout the book. Some entries
induce head-scratching. Van Tuinen
(p 409), for example, says that the
two major groups of living birds
(paleognaths and neognaths) separated
about 120 Ma, whereas
Benton and his coauthors list it at
about 66 Ma (“soft maximum”
dates are often ridiculously old and
can generally be disregarded). Here
is an example of where a character-based approach to fossils may
help constrain molecular estimates.
If one examines the fossil
record of birds about 120 Ma, and
even later in the Cretaceous, what
do we find? Well, the fossil birds
found in the Jehol Biota of China
(Early Cretaceous, about 125 Ma),
where the famous “feathered
dinosaurs” are also found, include
things like Confuciusornis that are
hardly advanced beyond
Archaeopteryx. Through the
Cretaceous we find thousands of
bird fossils, but they are all of primitive
toothed groups and
Enantiornithines, none of which is
regarded as close to Neornithes.
More importantly, the morphological
features that are in any way similar
to those of living birds do not
appear until the latest Cretaceous
(66–70 Ma). To accept the molecular
view of life, molecules are
doing the diverging, but this is very
seldom reflected in morphology.
Is this a reasonable view of life?
Space prohibits a review of the editors’
strange take on rates of diversification
of taxa, especially where
we have an actual fossil record that
is pretty reliable. Suffice it to say
that there is forty years of literature
on Phanerozoic diversity that
cannot be reduced to the unsatisfactory
alternatives of “dampened
exponential curve” or the “exponential model”. This book will be a
fabulous basis for advanced interdisciplinary
seminars, but I put the
accent on “interdisciplinary”.