toomey at museum.state.il.us
Tue Dec 5 18:49:44 EST 2000
> Fossils and Phenology in the Evolution of Ginkgo biloba,
> Rothwell,Holt, Ohio University
> Cladistics of the Spermatophyta, Brittonia, Loconte, Stevenson
> A reevaluation of seed plant phylogeny, Ann Missouri Bot Gard
> Lignophyte phylogeny and the evolution of spermatophytes, a numerical
> cladistic analysis, Syst Bot
> The Ginkgo Pages
Thank you for the references (although more complete citations to
make them easier to find would have been useful). I have
only been able to evaluate two of the four references that
you cite. However, neither of them provide support
for your contention that ginkgos might be the first
tree on earth.
The two that I can evaluate are as follows (references
Nixon KC. Crepet WL. Stevenson D. Friis EM.
A REEVALUATION OF SEED PLANT
Annals of the Missouri Botanical Garden. 81(3):484-533, 1994.
Seed plant phylogeny is
evaluated using a data set of 46 terminals (taxa) and 103
anatomical characters. Cladistic analyses using the criterion of
were performed on the complete data set as well as on subsets of the
e.g., excluding fossils and/or combining various complex taxa into
terminals. The results support the placement of the cycads as the
group of a monophyletic group that includes several fossil
''seed ferns'' as well as extant Ginkgo, conifers,
gnetopsids, and angiosperms. When fossils were included, Bennettitales
(cycadeoids) were part of an ''anthophyte'' clade that included
and angiosperms. Pentoxylon was a sister taxon to the core anthophyte
in some, but not all, of the most parsimonious trees. Caytonia was not
to be closely associated with the anthophyte clade, but instead was
associated as a sister taxon of the glossopterids, and these two taxa
consistently outside of the Ginkgo-conifer-anthophyte clade. In all
parsimonious trees for all analyses, Ephedra was to the outside of a
that included all angiosperm taxa, Gnetum, and Welwitschia, thus
the traditional gnetopsid clade paraphyletic. New information is
the morphology of Caytonia and some previous interpretations of
the caytonian ''cupule'' are rejected. The effects of sampling,
compartmentalization, and polymorphism are explored in these data,
how different results may be obtained when polymorphic or ''summary''
terminals are used. The need for more work on gnetopsids and fossil
suggested. [References: 234]
Rothwell GW. Serbet R.
LIGNOPHYTE PHYLOGENY AND THE EVOLUTION OF SPERMATOPHYTES - A
NUMERICAL CLADISTIC ANALYSIS
Systematic Botany. 19(3):443-482, 1994 Jul-Sep.
Phylogenetic relationships of 27 taxa of extant and extinct
assessed in a numerical cladistic analysis using 65 characters. The
yielded 12 most parsimonious trees, where fossil taxa were found to
crucial role in the reconstruction of spermatophyte
phylogeny. The results supported hypotheses of monophylesis
for spermatophytes, archaeopteridalean progymnosperms, medullosan
seed ferns, cordaiteans, anthophytes, and gnetophytes.
Progymnosperms, seed ferns s.l., and hydrasperman
seed ferns were found to be paraphyletic. A large clade that
previously has been referred to as ''platysperms'' or ''saccate
occurs in our results, but it is not characterized by either of these
features. The results reflect a hypothesized trend from large
leaves (the ''cycadophytic'' growth habit) to small leaves and
sporophylls (the ''coniferophytic'' growth habit) in several groups,
coniferophytes (including cordaites, conifers plus taxads, and
found to be polyphyletic. Surprisingly, inclusion of the Late
voltzialean conifer Emporia renders monophylesis for conifers
taxads) less parsimonious than polyphylesis under most sets of
Extinct taxa provided data to clarify the order of character
the evolution of spermatophytes. Their inclusion in the analysis also
clarify homologies among ovule enclosing structures that have been
''cupules.'' [References: 141]
These two references (although very good studies) are not capable of
actually providing evidence on whether ginkgos are the first tree
for two different reasons.
First, since they only deal with seed plants, they fail
to address whether any non-seed-bearing trees exist (the
answer is yes). So, even if ginkgos were the most primitive
seed plant, that would not necessarily make them the earliest
tree. To answer the question of whether ginkgos are the
most primitive trees, you must look at a phylogenetic analysis
that addresses all vascular plants (not just seed plants).
One recent such analysis is Doyle (1998). It finds scale
trees (Lycopsids) to be more primitive that gingkos.
Sphenopsids and ferns are also more primitive.
Within seed plants a variety of Paleozoic seed ferns
as well as Cordaitales may also be more primitive than
ginkgos. Each of these groups also has arborescent
forms that pre-date ginkgos (so the fossil record fails
to support the ginkgos as earliest tree scenario).
The Nixon and others paper you cite even indicates
several arborescent fossil taxa are potentially more
primitive than ginkgos.
The second reason these references are not actually
able to provide the data to support your statement
is more subtle. Both of these are phylgenetic studies
that provide information on the relationships of
the groups. They can indicate how primitive
the ginkgo group is; however, they do not indicate
that the earliest ginkgos are trees. Most of the
taxa have a variety of habits, so tree is not
necessarily the form in which they first show up.
As I noted in the first reason, the fossil record
shows a variety of trees that are earlier than
the earliest ginkgo.
I noted I have only been able to look at two of the references;
however, from the title, Cladistics of the Spermatophyta,
appears to suffer from the same problems as the two
I have discussed above in being able to provide evidence
on the claim at hand. Perhaps you can enlighten us with
better info on that one.
Reference cited (in addition to the ones cited by Cork)
Doyle JA, 1998, Phylogeny of vascular plants, Annual
Review of Ecology and Systematics. 29: 567-599.
Illinois State Museum
toomey at museum.state.il.us
> >CorK wrote:
> >> On 4 Dec 2000 21:39:40 GMT, una at mercury.cis.yale.edu (Una Smith)
> >> wrote:
> >> >
> >> >kwantenzap at xs4all.nl (CorK) writes:
> >> >
> >> >>It might well be the first tree on earth, you're not well informed.
> >> >
> >> >It is certainly not the first tree on earth. CorK, I suggest you go
> >> >*read a paleobotany textbook*; several good ones have already been
> >> >cited in this thread.
> >> I have.
> >CorK, since you indicate that you have read a paleobotany text,
> >perhaps you could tell us which one supports your claim that
> >the ginkgo "might well be the first tree on earth." As a
> >I would definitely support Una's statement that it is certainly
> >not the first tree on earth. In fact, it is not even close.
> >Trees predate the ginkgo line (let alone the genus Ginkgo), by something
> >like 50 million years.
> >Rick Toomey
> >Illinois State Museum
> >toomey at museum.state.il.us
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