| The
name Pliocene means "more recent", and this was the most recent
epoch of Tertiary period, lasting from about 5 to 2 million years ago.
Compared to previous epochs this was a relatively brief period, "only"
3 million years. During this time the world became much more like it is
today, with ice caps, modern mammals, relatively modern geography, and
the evolution of prehistoric man ("ape man").
Plate Tectonics and Geography
By the Pliocene the world was approaching that of today, and continents
had taken up their present-day positions. A shift in the Caribbean tectonic
plate, brought about the joining of North and South America, creating
a land bridge for mammals to migrate across. The Mediterranean sea (the
last remnant of the once mighty Tethys ocean) dried out, and was to
remain dry plains and grassland for several million years. During this
time, India collided with Asia and gave rise to the Himalayan Mountains,
the Himalayan uplift triggering a great global cooling (or accelerating
the already unfolding cooling process)
Climate
The Pliocene saw the continuation of the climatic cooling that had
began in the Miocene, with subtropical regions retreating equatorially,
the beginning of the large ice caps, especially in Antarctica, and
the northern hemisphere lands and ocean cooling likewise.
Antarctica was not yet completely
frozen. Throughout the Pliocene, Nothofagus remained common, as indicated
by pollen microfossils from this time.
In the northern hemisphere
there is a gradual southward migration of marine invertebrates. Reflecting
the cooling trend, Arctic species appear in Britain, and later, in the
Mediterranean. So much so that the horizon of a normal marine bed can
be determined by the percentage of species that (a) are extinct, (b)
survive in more northern latitudes, and (c) are today found in more
southern latitudes.
Pliocene Life
Pliocene Mollusks
The Pliocene saw an almost complete turnover of molluscan species
in numerous locations. This is not to say that extinction rates
approached 100% on the species level. In many cases, species ranges
simply flowed back and forth with shifting climate. This is particularly
noticeable in regions with long north-south coastlines, as in North
America. As the ice sheets spread, many of the endemic mollusks
of the California coast during the Zanclean were replaced in the
Gelasian by existing cold-adapted species from Canada and Alaska.
Some of the Zanclean Pacific coast species simply relocated to the
Gulf of California in the Gelasian, since the Gulf remained semitropical
throughout the Pliocene.

This is not to underestimate
the effects of the Ice Ages on molluscan diversity. However, an admittedly
cursory glance at the literature suggests that the net reductions in
molluscan diversity occurred later, during the Pleistocene. A recent,
very large-scale study of molluscan turnover by Todd et al. (2002) is
worth special emphasis. This work focuses on the Caribbean Basin from
the Late Miocene (12 Mya) to the Recent. Todd's group finds that the
driving factor in Plio-Pleistocene molluscan turnover was not temperature
change, but a steep reduction in nutrient levels associated with the
formation of the Isthmus of Panama and isolation of the Caribbean from
the Pacific. This was associated with a remarkable increase in molluscan
turnover throughout the Pliocene. However, there is no net decline in
diversity until the Pleistocene. Further, the decline in diversity is
explained almost entirely by a reduction in the number of gastropod
genera. Bivalve diversity remained largely constant. The authors explain
that reduction in overall nutrient levels favored reef-based filter-feeders
over gastropod carnivores.
The Todd study underlines
some points worth the emphasis. First, not everything in the Pliocene
can be explained by temperature reductions associated with the Ice Age.
The temperature in the Caribbean was constant and may actually have
increased slightly. Equally important were the changes in ocean circulation.
These changes may have had a role in creating the continental ice shelves
of the Gelasian. However, quite apart from arguably converting most
of Laurasia into hemispheric gelato, changes in ocean circulation had
very important effects in other ways.
Second, the Todd study sets
up an interesting line of speculation. Because it is one of a mere handful
of really large-scale, well-constrained diversity studies, we may have
confidence that our rampant speculations are at least grounded in reality.
The authors do not stress the point, but the delay between species turnover
and ecological collapse which appears in their data might be a general
phenomenon which would explain some of the inconsistent results in "Punctuated
Equilibrium" studies. Punctuated Equilibrium holds that evolution
happens in bursts, rather than at a continuous, rather constant clip.
Now, let us impose a few hypothetical changes in the isolation of the
Caribbean from the Pacific.
(1) Suppose that shortly
after the isthmus of Panama formed, some Pliocene Teddy Roosevelt had
gone down and dug the whole thing up. The brief isolation of the Caribbean
from the Pacific would then end, and Recent geologists would never learn
of the land bridge between North and South America. Three million years
later, we would observe, in the paleontological record, a rapid burst
of molluscan evolution, followed by a likely return to a more static
state, but with entirely new species, and without any obvious change
in the environment or ecosystem. In short, we'd see a punctuation episode.
(2) Now, take it a step further. Suppose Teddy had been delayed, perhaps
by a run-in with a bull moose, so that he didn't arrive in Panama until
after the collapse of the gastropods. However, he still digs up the
Isthmus so that we never learn about it. What we now observe is an "inexplicable"
mass extinction. (3) Finally, suppose that instead of Roosevelt, we
dispatched a horde of lawyers to Panama. The whole business would then
be tied up in litigation and nothing would get done properly. The land
bridge would be enjoined and never quite get finished, the digging up
process would bog down in permit applications, and everyone would ultimately
run out of money and patience. For the next 3 My, while the whole thing
slowly sleazed its way up to the Court of Geological Appeals, the Caribbean
and Pacific would be partially isolated -- perhaps only meeting at high
tide, or on alternate weekends and for three weeks in the summer. In
that case, it is likely that the gastropod clan would have time to adapt
to this change in its fortunes without ecological collapse. That is,
we would see a long, continuous process of more leisurely species turnover.
So, then, the question: are
mass extinction, punctuated equilibrium, and classical Darwinian gradualism
simply three points on a single spectrum?
As is all too often the case,
we have wandered so far off the intended path, that it would now be
hopeless to attempt any remarks concerning Pliocene mollusks. Accordingly
we will simply leave abruptly, salvaging whatever shards of dignity
may remain for some happier occasion
Pliocene Vertebrates
Terrestrial Ecosystems
The fauna of the Pliocene does not differ much from that of the Miocene,
although the period is regarded by many zoologists as the climax of
the Age of Mammals. This epoch is characterized by the appearance
of all of the presently existing orders and families, and many of
the existing genera of mammals.
Pliocene Vegetation was very
like today's. Grasslands replaced forests, so grazing mammals spread
at the expense of browsers. Cattle, sheep, antelopes, gazelles, and
other bovids reached their peak. North American mammals included horses,
camels, deer, pronghoms, peccaries, mastodonts, beavers, weasels, dogs,
and saber-toothed cats. Rhinoceroses and protoceratids died out in North
America. The one-toed horse appears for the first time.
Animal migrations
The Pliocene was a time of great migration, owing to the appearance
of new land bridges.
The North American three-toed
Hipparion horse crossed the Bering Straits land bridge and entered Asia
and Europe, while mastodonts entered the Americas from Asia.
During the late Pliocene,
about 3 million years ago, the isthmus of Panama ended South America's
isolation. The armadillo, ground sloth, opposums, and phorusrhacid birds
were among the animals that migrated North from South America. And dogs,
cats, bears, horses, mastodonts, and others animals invaded South America
from the north. This was catastrophic for some of the local animals,
especially the big marsupial carnivores. Even today more than half the
genera of South American mammals are descended from northern species.
Australia, still isolated,
saw rodents rafting in on mats of vegetation drifting south from Indonesia.
Hominid Evolution
Australopithecus robustus
In Africa the emerging
savanna grasslands and retreating forests caused some apes to come
down from the trees and take up life in the open, where they co-existed
with early elephants, antelopes, and other types of animals. An
erect posture was necessary for these vulnerable creatures to watch
for predators, which also freed the hands for the use of makeshift
tools (sticks etc). Thus the hominid lineage appeared in the rift
valleys of north-east Africa during Early Pliocene. As with the
bovids, the hominids underwent an evolutionary radiation, with a
number of lines of gracile and robust Australopithecines inhabiting
Ethiopia and Tanzania, and probably spreading throughout most of
Africa. The large-brained australopithecine Homo habilis continued
on into the Early Pleistocene, giving rise to Homo erectus , the
common ancestor of both Neanderthal and modern man during the late
Pleistocene.
The information
of this page came from http://palaeos.com/
For more
on the Cenozoic periods go to http://palaeos.com/Cenozoic/Cenozoic.htm
Page uploaded on WebDyer Site on 10 June 2004 ,
last modified 10 June 2004
text content © M. Alan Kazlev 1998-2002
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