Genome duplication in angiosperm evolution

I suspect that you will find interesting a proposal that angiosperm plants evolved on the Ontong-Java Plateau before the Cretaceous in http://charles_w.tripod.com/ontong.html and below; Also you may see an explanation for the boundaries of the temperate deciduous forest as a function of glaze ice storms in http://charles_w.tripod.com/glaze.html .

PERMIAN MONSOON EVOLUTION OF DECIDUOUS ANGIOSPERMS to
FURNISH CRETACEOUS GLAZE ICE TREES
by Charles Weber
ABSTRACT
Angiosperm deciduous dicotyledon trees have had much more success in surviving
in glaze ice areas than other types of trees because of less branch breakage. They
started to become established in North American subarctic regions in the Paleocene
and were fully established before the Eocene closed. They probably descended
largely from subtropical trees, those that had evolved hard wood because of
termites. I suspect that they may have evolved the precursor deciduous genes on a
now submerged Southwestern Pacific continent, the Ontong Java Plateau, probably
as early as the Permian, as a monsoon area adaptation. Their seeds were probably
carried to the mainland on the feet and in the crops of water birds.
DISCUSSION
Angiosperms are unique in having aromatic amino acids and hydrolyzable
tannins. I assume this helps toward insect resistance. Their carpels enclose their
ovules and pollen tube growth goes through sporophytic carpel tissue. All the
continents have been thoroughly explored and no large angiosperm assemblage has
been found anywhere before the Cretaceous. Evolution in tropical mountains
[Axelrod 1952, p34] is impossible. Such an enormous assemblage of mono- and
dicotyledons would have been absolutely certain to have left many obvious lowland
fossils in view of their considerable success not much later. Every other plant group
can be traced back at least to the Permian and most go back to the Devonian
[Stewart p128,169,212,313,348]. Molecular evidence shows the monocotyledons
separating from the dicotyledons 320 million years ago in lower Carboniferous
[Martin et al, 1989] or from phytochrome gene about 260 million (lower Triassic)
[Kolukisaoghi, et al, 1995]. Molecular analysis of other major divergences are 200
million years or more. There are leaves and flowers in Triassic Virginia [Cornet,
1993] and also pollen [Cornet, 1989] resembling angiosperms [Cornet, 1989] as
well as aquatic fossils with fruit from late Jurassic in Central Asia [Sun, et al, 1998;
Sun, et al, 2002] so it is plausible that angiosperm prototypes survived in small
numbers as late as the Triassic or later on the continents. Isocarbinorinol, which
only angiosperms have, has been found in the Permian [Haike, et al 1995] so this
possibility is reinforced. There is even a report of pollen resembling angiosperm
pollen from the Jurassic North Sea [Abbink, 1998]. This pollen of a water plant
could conceivably be an early migrant however. A tropical island that could field so
many tropical families would have to be large. A string of offshore volcanic islands
or even peaks of a mountain range would be off by at least two orders of
magnitude. The only unexplored areas that I know of are in the western Pacific.
Elevation of the Ontong Java Plateau would create a fair amount of tropical land, the
size of Alaska or a major part of Australia, the largest oceanic plateau. Some of that
area could have been monsoon climate that also can favor a deciduous habit.
Seismic evidence indicates that this rise has continental characteristics with a crust
over 35 km thick [Furomoto, et al, 1976]. There is no evidence of flexure under the
rise [Hussong, et al, 1979] so the rise must go back to the very beginnings of earth
history. If we drill through the basalt someday we may find coal or pollen
underneath the basalt and thus possibly throw light on this. This has not been done
to date [Kroenke, et al, 1990]. There also should be angiosperm pollen in mid
Cretaceous sediments near the sea mounts between the Ontong Java Plateau and
Korea, where the Angiosperms first appeared, if this proposal is correct.
New Caledonia has many very primitive angiosperms [Carlquist, 1965 p74].
New Caledonia and Lord Howe Rise near New Zealand are possibilities as survival
centers since they were probably connected together at one time based on similar
creatures which cross oceans with difficulty [Paramonov, 1963]. A string of off
shore islands and sea mounts as the mini continent sank to provide magma for the
nearby sea mounts in the beginning of the Aptian [Rohl, 1996] (the Ontong-Java
plateau had shallow water in early Cretaceous [Nur and Ben-Avraham, p3644] )
could have provided a holding area to save an already evolved flora from extinction.
Indeed, some very primitive angiosperms have survived until modern times on New
Caledonia and other Pacific Islands [Melville, 1966]. That no mammals or snakes
live on New Caledonia is circumstantial evidence that it was not attached to the
mainland recently [Carlquist, 1965 p74], and probably never. That does not rule out
the possibility that at some time in the past it furnished plants to Lord Howe Rise
across narrow water gaps. Pollen has been discovered from the Permian, Triassic,
and Jurassic on New Caledonia [Dejersey & Grantmackie, 1989]. None of this
pollen resembles angiosperm pollen [Raine and de Jersey, private communication],
so New Caledonia is virtually ruled out except as a center of survival. Long water
gaps were not too likely possible since there were no birds then and no spore
reproduction in angiosperms, so primitive angiosperms could have been almost as
poor as conifers are today at crossing oceans. However, the modern angiosperms
are fair to excellent at jumping ocean gaps today [Carlquist, 1965 p35- 38] so the
islands near here could have contributed both to further evolution and conservation
after seed eating or water birds arrived. That seed eating birds were present in early
Cretaceous has always been plausible, but now the discovery of a seed eating fossil
bird from then recently has made it certain [Zhou and Zhang, 2002]. At the same
time, angiosperms could have been unable to bridge a larger gap to the continents
until early Cretaceous when the gaps narrowed because of the rise of sea mounts
between the Ontong Java Plateau and Korea [Larson, 1991; Rohl, 1996] and
efficient birds appeared. The absence of mammals on all the islands just mentioned
imply a separation from the mainland going back possibly to the Triassic or further.
The fact that angiosperms lacked an efficient means of dispersal by mammals in the
first half of the Cretaceous [Schuster, 1976, p52, 64] is circumstantial evidence that
they came from an island continent. Fruits and barbs are late in angiosperms.
It must seem obvious that the above discussion is at odds with some of current
plate tectonic theory. However, shallow earthquakes under the longest ridge-ridge
transform faults make it seem to me [Weber, 1981;
http;//members.tripod.com/~charles_W/ridge.html ] that these islands were always
where they are now.
That deciduous evolution was at least partly subtropical in the Cretaceous,
however, is almost certain. The first plants were herbaceous, insect pollinated,
[Stewart, 1983, p370], small, arid habitat, streamside shrubs, [Hickey and Doyle,
1977] their seeds possibly carried in the crops or on the feet of waterfowl.
Monocots came much later followed by wind pollination [Stewart, 1983 p373].
Further migrations had all the earmarks of intermittent arrivals. It was not until late in
the Cretaceous that angiosperms moved toward the poles [Crane and Lidgard,
1989; Parish and Spicer, 1988] aided by a warming trend, probably by evolution in
subtropical genera on the fringes. The late Cretaceous was a rather warm time
[Jenkyns, et al, 2004].
The colonization of the Northern Hemisphere by the hardwoods may have been
considerably aided by the presence of Neotermes. This is a termite genus which
was likely to have been able to eat live wood as early as the lower Cretaceous
[Weber, 1993]. An ability to girdle taproots may have assisted it then and does
today. There were no woodpeckers or Ponerine ants in monsoon areas, or other
effective predators. Even if there had been woodpeckers, they would not have
touched Neotermes if they had been like modern woodpeckers that do not
[Kalshoven]. Strong wood with living tissue on the perimeter would have
preadapted the above trees to a trend that was already visible in other early
Cretaceous trees. Neotermes can not kill a hardwood unless the hardwood is also
hard hit by drought or disease [Hill, 1921]. The ability of Neotermes to migrate on
floating logs makes its early genesis less than certain, but its primitiveness [Snyder,
1949] makes it plausible. The later rise of the angiosperms to 85% of the fossil
species [Axelrod, 1966] {but not necessarily 85% of the area covered} may have
been assisted by the appearance of Coptotermes in the rain forest, which insect also
uses live wood, toward the end of the Cretaceous in the Northern Hemisphere
tropics. Most termite species can utilize susceptible live wood, but the appearance
of angiosperms came almost certainly at least 50 million years after termites
appeared as fossils so the trees must surely have evolved some chemical defenses
against termites or their protozoa by the time angiosperms appeared as fossils since
Neotermes can spread on floating logs. It is possible that Coptotermes ability to
secrete its own cellulase [Hogan, et al, 1988] may make Coptotermes able to utilize
trees protected only by poisons against microorganisms. Neotermes ability to
ignore the Nasutitermes alarm pheromone secreted by pine trees (alpha & Beta
pinenes) which prevent Nasutitermes from living under pine trees [Lee and Wood,
1971, p7] must have assisted Neotermes in clearing away pine trees, at least when
Nasutitermes reached North America, since Nasutitermes probably evolved during
the Cretaceous in South America and presumably Nasutitermes would have
competed indirectly somewhat otherwise. Coptotermes presence by late Cretaceous
is backed by present continental distribution but not by fossils yet. Its superior
defense, utilizing a poisonous sticky secretion, must surely have made it successful
even in rain forests, since there were probably no Dorylene ants in mid Cretaceous
in the Northern Hemisphere (there is a good chance they were already in South
America), and Ponerine ants probably did not then hunt in packs based on present
day distribution.
Glaze ice must have existed in the late Cretaceous because there was deciduous
vegetation in northwestern North America [Wolfe, 1986]. Also rapid changes in sea
level even before that time imply the presence of glaciers somewhere [Stoll and
Schrag, 1996].
Apparently the Carboniferous or Permian monsoon genes which preadapted
angiosperms to glaze ice forests [http://glaze.freeyellow.com/index.html ] were already in
place. Numerous tropical and subtropical genera moved into a glaze ice climate
zone, which today is centered in Northern Illinois, during the Tertiary starting in late
Cretaceous [Spicer and Parish, 1986; Spicer and Chapman, 1990] by displacing
deciduous Metasequoia [Wolfe, 1987, p219] and other deciduous conifers [Wolfe,
1986]. The subtropical hardwood genera predominated, so that evergreen maples
{Acer}, subtropical deciduous oaks {Quercus}, beeches {Fagus}, Sycamores
{Platanus}, and subtropical maples {Acer}, as well as trees whose climate
forebears were not specified by Axelrod [1966] such as hickories {Carya},
chestnuts {Castanea}, and elms {Ulmus} took over the canopy in the fertile heart
of the glaze ice zone, leaving tropical and other subtropical genera to fill in the
chinks, the understory, sandy or acid soils, the periphery, the disturbed areas, and
the swamps. Almost all of the above hardwoods were or became deciduous.
There must have been a zone of glaze ice in Alaska if this hypothesis is valid
because several species of southern Appalachian plants are also known in Asia
[Braun, 1972 pp515-517, 460]. Therefore it is conceivable that at least a narrow
band of glaze ice existed across the Bering Straits bridge, Pliocene or earlier. The
Paleocene vegetation at 66 degrees north was largely deciduous [Wolfe, 1987],
which lends plausibility to the above statement. If this hypothesis is valid, it explains
why the extravagant deciduous trees which lose all their leaves annually, hang their
soft edible leaves out within easy reach of sucking and other insects and
vertebrates, and show extreme difficulty when competing with pines and sequoias
over vast cool areas, nevertheless make almost monolithic stands in a
gerrymandered area which cuts across isopleths of humidity, temperature, rainfall,
light duration, light angular incidence, soil, fire, and nutrient status in temperate
regions.
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