On 10/8/14, 5:56 AM,
nyi...@bellsouth.net wrote:
> On Tuesday, October 7, 2014 11:26:19 PM UTC-4, John Harshman wrote:
>> On 10/7/14, 7:43 PM,
nyi...@bellsouth.net wrote:
>
>>> One way of minimizing this contrast is to claim that the concept of "phylum"
>>> is an artificial one, but it has been used for well over a century to
>>> be roughly synonymous with "unique body plans." So it is an indicator
>>> of disparity, the idea being that animals within a phylum are less
>>> different in some fundamental way from each other than any in one phylum
>>> are from any in another phylum. And so perhaps the real question about
>>> the Cambrian explosion is this: why did so much disparity blossom in so
>>> short a time, while later in earth history it happened on a smaller scale
>>> in a much longer time?
>
>> Back when I paid attention to the literature on this, there were three
>> main ideas.
>
> How long ago was that? Can you recall any websites (or at least
> articles or books) where the following ideas were advanced?
Here is something I wrote for a seminar on the subject back in the
mid-'90s. The references should get you into the literature.
The Cambrian Explosion
What do we see?
The first unquestioned metazoans appear in the late Riphean or early
Vendian, loosely termed the Ediacaran fauna. They are all soft, and are
mostly preserved as molds in coarse sandstones. There are also trace
fossils, all (?) surface trails. Near the end of the Ediacaran, the
Ediacaran fauna fades out, trace fossils diversify, and the first
"small, shelly fossils" appear. The Tommotian is a period of gradual
increase in diversity of the shelly types, including the first examples
of modern phyla, and of trace fossils. The first trilobites appear in
the Atdabanian, as do (?) the first post-Ediacaran unmineralized
metazoans. All modern phyla are known by the end of the Cambrian, all
classes (of marine invertebrates) by the end of the Ordovician (not
counting those without fossil records).
What needs explanation?
Why didn't this happen earlier? Why not later? Why so fast? Why only
once? The fourth question is the only one I intend to worry about here:
why did almost all the animal phyla (apparently) arise in the Cambrian,
and none since then? (Say, what about plants? What about land animals?)
Explosion? What Explosion?
Maybe the supposed explosion is due to taxonomic or taphonomic effects.
Taxonomic: Raup has a complicated simulation, but I think it boils down
to the idea that in a nested hierarchy, deep nodes have to come before
shallow nodes. If higher taxa are not defined cladistically, is this
relevant?
Conway Morris (1985, mostly) has another idea. Maybe these perceived
radical differences in bauplan are artifacts of the big gaps between
modern groups; a Cambrian systematist would see only low-level taxa
where we see phyla and classes. Briggs & Fortey suggest something
similar in reference to the arthropods.
Taphonomic: Do the various groups have invisible Precambrian histories?
Valentine & Erwin examine various ways for this to happen: ancestors
were soft & squishy, teeny, rare or localized, lived in unpreserved
environments; the fossil record sucks. They reject all of them.
Are we looking in the right place for Precambrian ancestors? Where are
the animals that made Precambrian trace fossils? Were Ediacaran fossils
especially preservable or were the rules of preservation different then?
See Seilacher, Fedonkin.
Butterfield examines in detail the factors involved in preservation of
the Burgess Shale. Animals are preserved as films of original carbon. An
extracellular cuticle with various means of keeping its proteins from
attack (crosslinks, e. g.) is almost universal. Since animals carry the
means of their own destruction within their guts (proteolytic enzymes
and intestinal biota), filling of coelomic spaces with fine clay both
adsorbs enzymes and blocks spread of decomposers. How about a late
evolution of cuticle? Of coeloms? (But what about body plans that just
won't work at all without rigid parts?)
The Empty Barrel
You know the story: when the starting gun goes off, everyone races to
fill vacant ways of life. With plenty of room and no competition,
selection is relaxed and experiments are possible. When the barrel is
full, there's no room for experiment. In the end Permian, species
diversity crashed but ecological diversity was still high: the barrel
was still mostly full, thus no room for new phyla (Erwin et al.) How
does this scenario cover the invasion of the land?
McMenamin's variant makes the origin of predation the cause, with no
required relaxation of selection (quite the opposite). Once the noise
died down, it would take a new stimulus to set things going again, which
has not happened yet.
Kauffman sort of fits here, although he requires no biotic
intereactions. He envisages adaptive landscapes, with species starting
in valleys. At the beginning, a big jump is as likely to improve fitness
as a small jump, but this ends soon. Why are species starting in valleys?
Loose Genes
The basic loose gene theory waves its arms and talks about
canalization. Epistatic effects were less common then. (Is this
possible?) Maybe epigenetic homeostatic mechanisms (say it three times
fast) weren't as fully evolved as they are now. (Would this result in
greater embryonic mortality? Is there a way to look for that?) Jablonski
& Bottjer suggest a test by examining deviations from bilateral symmetry.
Jacobs at least tries to elaborate his mechanism enough to be testable.
In arthropods, in some systems, at least, serial genes divide the
developing body into identical segments, while selector genes, with
regional patterns of expression, differentiate the segments. He supposes
this to be the ancestral development pattern
of coelomates, which are thus primitively segmented. He divides phyla
into two groups: segmented taxa which have presumably retained the
ancestral system, and non-segmented taxa which have not. In segmented
types, selector genes were once closely linked and were open to all
sorts of simple rearrangements with big phenotypic effects. Now they are
scattered over the genome and big changes are no longer easy. Ordinal
originations of segmented animals show such a historical pattern, while
non-segmented animals do not.
Problems? Patel et al. come to a different conclusion on ancestral
states: the ancestor had a serially ordered central nervous system, but
body segmentation evolved separately in arthropoda, annelida, and
chordata. Jacob explains why segmented animals used to have loose genes,
but why should unsegmented animals still have loose genes just because
regulation of development is conceptually more complicated? And of
course this pattern only explains the ordinal level, not class or
phylum. How well does Lake's phylogeny match Jacob's theories? Patel et al.?
The Cambrian Explosion: Bibliography
Anderson, M. M. and S. Conway Morris. 1982. A review, with descriptions
of four unusual forms, of the soft-bodied fauna of the Conception and
St. John's Groups (late-Precambrian), Avalon Peninsula, Newfoundland.
Third North American Paleontological Convention, Proceedings vol. I, 1-8.
Possibly the oldest known diverse fauna, perhaps as old as late Riphean,
and grossly resembling the Ediacaran fauna.
Briggs, D. E. G. and R. A. Fortey. 1989. The early radiation and
relationships of the major arthropod groups. Science 246:241-243.
Cladistic analysis of Burgess Shale and other arthropods shows
trilobites as highly derived, crustaceans as possibly paraphyletic, and
disparity within Burgess arthropods not especially high.
Butterfield, N. J. 1990a. Organic preservation of non-mineralizing
organisms and the taphonomy of the Burgess Shale. Paleobiology 16:272-286.
Talks about reasons for preservation of "soft-bodied" animals, including
a decay-resistant, extracellular cuticle and interior coelomic spaces
which can be permeated with clay particles. To add to hypotheses of
multiple, simultaneous skeletonization events, can we try
cuticle-ization and coelomization?
Butterfield, N. J. 1990b. A reassessment of the enigmatic Burgess Shale
fossil Wiwaxia corrugata (Matthew) and its relationship to the
polychaete Canadia spinosa Walcott. Paleobiology 16:287-303.
Wiwaxia is a polychaete, reducing the number of funny phyla by one.
Chen, J. 1988. Precambrian metazoans of the Huai River drainage area
(Anhui, E. China): their taphonomic and ecological evidence.
Senckenbergiana Lethaea 69:189-215.
Another candidate for oldest metazoans, this time annulated, coelomate,
burrowing worms, dated at 740ma. Big implications if so. But see Conway
Morris 1990.
Conway Morris, S. 1985. Cambrian Lagerstatten: their distribution and
significance. Philosophical Transactions of the Royal Society of London
B311 ꌴ9-65.
Survey of Cambrian soft-bodied fossil localities. Considers the
explosion of phyla as an artifact of hindsight: we see the Cambrian
world through a filter of living groups.
-----------. 1989a. Burgess Shale faunas and the Cambrian explosion.
Science 246:339-346.
Description of the Burgess Shale fauna. No useful input on the
uniqueness of the Cambrian explosion.
-----------. 1989b. The persistence of Burgess Shale-type faunas:
implications for the evolution of deeper-water faunas. Transactions of
the Royal Society of Edinburgh: Earth Sciences 80:271-283.
At least some of the Burgess genera have very long duration (stasis?),
some from ?Tommotian (more likely early Atdabanian -- see Dzik and
Lendzion), arguing for a rapid Cambrian explosion. (The main subject of
the paper is some gibberish about taxa originating in shallow shelf
environments.)
-----------. 1990. Late Precambrian and Cambrian soft-bodied faunas.
Annual Review of Earth and Planetary Sciences 18:101-122.
A short review of Ediacaran and Burgess fossils.
Donovan, S. K. 1987. The fit of the continents in the late Precambrian.
Nature 327:139-141.
Just thought you might wonder what the world looked like; three theories
evaluated from a biogeographic perspective.
Dzik, J. and K. Lendzion. 1988. The oldest arthropods of the East
European Platform. Lethaia 21:29-38.
A third species of anomalocarid and a naraoid trilobite from the
?Atdabanian are the earliest Burgess-like fossils known.
Erwin, D. H., J. W. Valentine, and J. J. Sepkoski, Jr. 1986. A
comparative study of diversification events: the early Paleozoic versus
the Mesozoic. Evolution 41:1177-1186.
Documents the explosion and gives a classic statement of the empty
barrel theory.
Fedonkin, M. A. 1985. Precambrian metazoans: the problems of
preservation, systematics and evolution. Philosophical Transactions of
the Royal Society of London B311:27-45.
An interesting view of Ediacaran taphonomy. That's the good part. The
rest is weird morphological evolutionary models based on superficial
resemblances. Annelida from Cnidaria in two easy steps?
Field, K. G., G. J. Olsen, D. J. Lane, S. J. Giovannoni, M. T. Ghiselin,
E. C. Raff, N. R. Pace, and R. A. Raff. 1988. Molecular phylogeny of the
animal kingdom. Science 239:748-753.
18s RNA-derived phylogeny presents several mutually inconsistent trees
containing different combinations of taxa. A consensus tree of this
unintended jackknifing would show very little structure at the phylum
level. Maybe this suggests the coelomate radiation was very fast
(Cambrian explosion?). Or maybe it's just garbage.
Gehling, J. G. 1987. Earliest known echinoderm -- a new Ediacaran fossil
from the Pound Subgroup of South Australia. Alcheringa 11:337-345.
This replacement for Tribrachidium at least has five-fold symmetry. This
is here because the accompanying photos show how hard it is to tell much
from Ediacaran fossils.
Glaessner, M. F. 1984. The dawn of animal life: a biohistorical study.
Cambridge: Cambridge University Press.
A paleontologist/morphologist of the old school interprets Ediacaran
life, assigning everything to modern groups. Could be, but it sounds
like a suspicious coincidence to me.
Gould, S. J. 1989. Wonderful Life: The Burgess Shale and the Nature of
History. New York: W. W. Norton & Company.
Plenty of drama and great pictures. The explosion itself is not central
to his personal ax-grinding, but he has a nice summary on pp. 228-233.
Gould could perhaps use a short course in phylogenetic systematics.
Hoffman, A. and M. H. Nitecki (eds.). 1986. Problematic Fossil Taxa. New
York: Oxford University Press.
Authors tackle various weird groups, most of them relevant to the
Cambrian explosion. Lots of good photos and reconstructions. Bengston's
introduction has an interesting discussion of what we mean by "phylum".
Jacobs, D. K. 1990. Selector genes and the Cambrian radiation of
Bilateria. Proceedings of the National Academy of Sciences 87:4406-4410.
A previously unrecognized pattern in the evolution of novelties and a
"loose genes" explanation to account for it. This may be testable.
Kauffman, S. A. 1989. Cambrian explosion and Permian quiescence:
implications of rugged fitness landscapes. Evolutionary Ecology 3:274-281.
A third hypothesis, probably not testable. On an adaptive landscape with
many local maxima, assuming a species begins with low fitness, big
mutations are as likely to improve fitness as little ones. As a species
finds higher peaks, big mutations no longer work well. Thus phylum-sized
changes go away. There is a vague implication that the landscape gets
smoother too, which also makes big mutations unprofitable. But why
assume low fitness at the start?
Lake, J. A. 1990. Origin of the Metazoa. Proceedings of the National
Academy of Sciences 87:763-766.
Reanalysis of Field et al. using Lake's own parsimony method makes
arthropods paraphyletic (?) to a big group including Annelida, Mollusca,
and Brachiopoda. Some basal branch lengths are long, implying
considerable undiscovered Precambrian history of coelomates. Weird
stuff, but food for thought.
Matthews, S. C. and V. V. Missarzhevsky. 1975. Small shelly fossils of
late Precambrian and early Cambrian age: a review of recent work.
Journal of the Geological Society of London 131:289-304.
I picked this one because it has photos of most of the "small, shelly
fossils" you hear so much about. Plate 4 figs. 2, 5, 8 is Microdictyon,
unnamed until 1981.
McMenamin, M. A. S. 1989. The origins and radiation of the early
Metazoa. In: K. C. Allen and D. E. G. Briggs, eds. Evolution and the
fossil record. London: Belhaven Press, 73-98.
The Reader's Digest version of his book, below.
McMenamin, M. A. S. and D. L. S. McMenamin. 1990. The Emergence of
Animals: the Cambrian Breakthrough. New York: Columbia University Press.
A good semi-popular account of metazoan history through the Cambrian,
including his variation on the ecological explanation for the explosion:
the invention of predation caused a revolution, devices for avoiding
predators becoming exapted for new ways of life. Things settled down
soon, and permanently, for reasons that aren't clear to me, but the
implication is that only in periods of ecological chaos are there
opportunities for new body plans.
Patel, N. H., E. Martin-Blanco, K. G. Coleman, S. J. Poole, M. C. Ellis,
T. B. Kornberg, and C. S. Goodman. 1989. Expression of engrailed
proteins in arthropods, annelids, and chordates. Cell 58:955-968.
This is a whole new world. I picked this as representative of numerous
papers cited by Jacobs because it covers the widest ground and draws
some phylogenetic conclusions for itself, which conflict somewhat with
Jacobs. Note especially what the annelids are doing. For an interesting
time, try to reconcile this paper with Lake.
Raup, D. A. 1983. On the early origins of major biologic groups.
Paleobiology 9:107-115.
Divergence time between >90% of major groups is predicted to be >500ma,
given the model of stochastically constant evolutionary rates. If we're
talking about evolutionary novelties rather than branching pattern,
divergence time is probably not a relevant measure.
Seilacher, A. 1984. Late Precambrian and early Cambrian Metazoa:
preservational or real extinctions? In: H. D. Holland and A. F.
Trendall, eds., Patterns of Change in Earth Evolution, Berlin:
Springer-Verlag.
Discusses the taphonomy of Ediacaran organisms, coming to the conclusion
developed in more detail in the next paper.
-----------. 1989. Vendozoa: organismic construction in the
Proterozoic biosphere. Lethaia 22:229-239.
Interpretation of the Ediacaran fauna as the separate phylum Vendozoa,
making the Cambrian explosion more explosive than ever.
Valentine, J. W. and D. H. Erwin. 1987. Interpreting great developmental
experiments: the fossil record. In: R. A. Raff and E. C. Raff, eds.,
Development as an Evolutionary Process. New York: A. R. Liss, 71-108.
Contains an analysis of various processes which might produce an
apparent (or real) Cambrian explosion and concludes that something weird
was going on, but makes no firm statement about what, finding all
current theories inadequate. Joel Cracraft memorial quibble: I wish they
wouldn't talk about "ancestral clades".