William H. Calvin, A Brain for All Seasons: Human Evolution and Abrupt Climate Change (University of Chicago Press, 2002). See also http://WilliamCalvin.com/BrainForAllSeasons/Equator.htm.
ISBN 0-226-09201-1 (cloth) GN21.xxx0
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William H. Calvin
University of Washington
modern rain forest stretches from the Atlantic to eastern [Congo],
which is closer to the Indian Ocean. During glacial maxima, it seems,
the rain forest shrank to three small patches, one near each end of its
present extent and the third in between, in southern Nigeria and
These three oases sometimes added up to no more than about 20
percent of the present extent of the African rain forest.
Steven M. Stanley, Children of the Ice Age, 1996
above the equator gives a view of the right bank and the left bank of
the Congo River, the dividing line between our cousins, the chimp and
the bonobo, starting about 2.5 million years ago – about the same
time that our earliest Homo ancestors split off from the bipedal
apes, probably somewhere east or south of here.
Lots of swamp forest down below, which is where the last bonobos
live. (They’re called
the “Left Bank Chimps” for other reasons, as well.)
They were trapped there by climate change, in one of the refugia
from the cooling and drying episodes.
The bonobos in the other refugia probably died out.
Certainly in the time since the orangutans split off the great
ape tree at about 12 million years ago, climate has changed.
Globally it cooled – and, in East Africa, the effects were
even more pronounced because the highlands of Kenya and Ethiopia were
pushed up like a blister, so that now Nairobi is slightly more elevated
than mile-high Denver. The
largest cities of a number of eastern and southern African countries
are equally elevated (which is fortunate, as it gets them up above the
lower-lying malaria zone).
Temperatures decrease about 6°C for every thousand
meter rise, everything else being equal, so Nairobi ought to be about
10°C cooler than Kenya’s Indian Ocean coastline – and
up here in the stratosphere at 10 km, the air ought to be about 60°C
cooler than down on the ground. The
two East African blisters thinned out the forest and created more open
woodland and even savanna (grasslands with the occasional tree).
Something similar probably happened in South Africa, where the
first australopithecine skull was found.
At the same time that things cooled, climate became much more
variable – so the averages don’t tell you the whole story.
(Averages are just devices to keep us from thinking more
deeply.) Just as
variability in populations is the key to thinking about how species can
change, so variability in climate (especially the fire-prone droughts)
is the key to thinking about what speeds up species change.
don’t know much about how the brain reorganized during the
last five million years, though we do know one tantalizing feature of
average size. Unlike
upright posture which was pretty well established before four million
years ago, brain size didn’t increase much until the Homo
lineage was spun off from the australopithecine lineage about 2.4
million years ago. Since
then, the brain has increased about three-fold in volume (and more like
four-fold in neocortical surface area, from more infolding).
Furthermore, sexual dimorphism decreases in the Homo
lineage; instead of australopith males being twice as large as females
(usually a sign of males fighting one another over access to females),
the Homo erectus males revert to being only 20 percent
larger than females, a pattern much like our chimpanzee cousins.
And as I earlier mentioned, the archaeologists say that
split-cobble toolmaking also started up about 2.5 million years ago.
Why was there all this prehuman action starting about 2.5
million years ago?
The first clue is that it isn’t just the hominid lineage that
spun off new species about then. That’s
also when the ancestral Pan lineage split into what is today the
bonobo and the chimp. The
bonobos are confined to the left bank of the Congo River, and now
limited to the forest immediately below me, here at Latitude Zero.
The various subspecies of the common chimpanzee extend across
equatorial Africa from the Rift Valley into westernmost Africa.
About 2.5 million years ago is also about when, among the lesser
apes in southeast Asia, the gibbons spun off the siamangs.
And numerous other major mammalian groups such as pigs and
antelopes also show a lot of speciation happening about then, best
documented in Africa. This
speciation fest, by itself, suggested big climate change between 3 and
2 million years ago.
The second clue is that Northern Hemisphere glaciation
intensified between 3.1 and 2.5 million years ago, thanks to all the
moisture delivered to the far north via evaporation from a more
vigorous Gulf Stream. Ice
sheets eventually built up to the height of mountain ranges over Canada
and Scandinavia. The ice mountains tended to melt off every 40,000 to 100,000
years, only to rebuild again. Yet
the site of the hominid speciation action was likely in the tropics,
probably somewhere here in Africa.
And Africa wasn’t icy back then – a little cooler (3-5°C),
but not the sort of thing that would keep the tropics from being nice
and warm in most places. Most
importantly, Africa was much drier (Lake Victoria, one of Africa’s
largest lakes, dried up during the last ice age).
So what do the Ice Ages have to do with stimulating all this
evolutionary action among the mammals – and particularly among our
African ancestors? (Think
drought, not cold.) What
might it have to do with the items on my little chunnel-train list of
the big-time augmentations of chimpanzeelike behaviors?
First let me explain some of the standard story about climate
change and population fluctuations.
Then I’ll try to show what abrupt climate change adds to the
story, which is a much less settled question, not yet in the textbooks.
saw that climate had repeatedly changed but, unlike others
before him, he successfully figured out a mechanism whereby animal
species could change with it, to adapt body and behavior to the new
climate regime. Just spawn
a lot of variations in each generation and, given the high mortality
among the young, only those variants better adapted to the current
environment will survive long enough to reach reproductive age.
And those lucky variants will spawn additional variations around
their body-and-behavior traits, to further explore “fits” to the
environment’s opportunities and perils.
Those variants better suited to some other climate simply tend
not to grow up and reproduce.
But note that this need not be sustainable change.
When the climate changes back to the original, the adaptations
can track it back again. (Remind
me later to explain how speciation can ratchet the adaptation, so it
doesn’t drift back so easily). Furthermore,
adaptations may mostly happen when there is no other choice.
At least on some time scales, climate’s influence needs to be
viewed with some skepticism, as most species react to cooling and
drying episodes by moving elsewhere, places where their suite of
adaptations still works.
Well, moving is something of an euphuism; if there are
regional subpopulations, some of them may die out while others
continue. With serious
climate change, this may leave only a few subpopulations in refugia,
places where the species still has all of the essentials for making a
living and reproducing. Let climate improve, and they will “expand their range”
to live in more places, with refugia pioneers rediscovering those old
places where the species once thrived.
Population size is always fluctuating like this.
A shrink-and-expand cycle produces more evolutionary change than
adaptations-in-place, as I mentioned earlier, but other factors that
truly fragment the central population may prove even more important in
transforming the species, particularly (as I’ll mention in a minute)
because of the chance aggregations that occur when things fragment.
Population fragmentations are what happens
when a lake almost dries up. As
the water level drops, you get a series of small ponds and puddles, in
which life continues – but there’s now a lot of inbreeding because
they are trapped and cannot circulate.
There may be some selection for living in the increasingly salty
ponds. Most little ponds
dry up completely, and the life in them doesn’t contribute to what
happens later (there are some exceptions, animals whose
lay-them-and-leave-them young can survive desiccation).
If only the population in one pond survives and then re-expands,
we see a classical “population bottleneck“ where the re-enlarged
population is comprised of only closely-related individuals.
Note that much of the pre-existing variety may vanish, even
though little natural selection affected the survivors directly (it
just eliminated much of their competition by chance).
Refugia are common on land, too, and land animals can be
similarly restricted to inbreeding for awhile, with a great reduction
in genetic variety because of sheer chance.
Cheetahs, all very similar genetically, likely re-expanded from
one such small surviving population.
This means that natural selection no longer has much variation
to operate on, preventing evolution until mutations and cross-over
breaks eventually generate some new variation on which recombination
But more often, multiple ponds survive the downsizing and
fragmentation. When the
old lake refills with the rains, multiple small groups form the basis
of the re-enlarged animal population.
Each group may have survived for a different reason, some
developing adaptations but most not. It is presumably only when the land refugia are also under
stress, when they too become excessively cool or dry or dusty, that
selection can efficiently operate to improve thermal regulation or
kidneys or noses. Or to
select for rare abilities that, for once, make a difference.
This seems fundamentally different from “who survives”
during ordinary population contractions into unstressed refugia.
And there is really nothing to suggest that it was all that cold
in tropical refugia for our ancestors.
Drought, however, is another matter, as is an ecosystem that
fire has severely disrupted. Cooling
is just the easiest thing to measure in reconstructing paleoclimate,
and not necessarily the most relevant thing to survive and thrive.
Lake Victoria, right on the equator over in East Africa, dried
up during the last ice age and abruptly refilled about 15,000 years
ago. The cichlid fish in
the East African lakes split into many new species about then.
Selection during downsizing isn’t the only way that evolution
operates. There are also
opportunities to be exploited when conditions improve.
And for an ape-like creature already adapted to making a living
on the savanna, the opportunities were of boom-time proportions.
changing climate drives different populations apart and
brings them together again. This
could have facilitated speciation.
Richard Potts, 2001
later part of the Pleistocene had been a period of extreme fluctuation
in climate. Vegetation
zones had moved north and south, up-mountain and down.
Extensive woodlands had been
fragmented by invading steppe or savannas, and had been rejoined
as forests returned. Glaciers and
harsh periglacial climates made vast areas of northern Eurasia periodically
uninhabitable by hominids, presumably spurring
major migrations and causing local extinctions.
Sea levels had risen and fallen, alternately
creating islands and land bridges.
Perhaps no period in the history of the globe had been more
conducive to the emergence of new species and to competition
between related species newly in contact – in other words, to
evolutionary change. And
the variety seen among later Pleistocene hominid fossils was, in fact,
exactly the kind of thing one might expect to find under those
- Ian Tattersall, 1995
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