Chapter 4
Lost Serengetis 1: Europe, with a
brief look at Asia
I’m going to talk, presently, about the
rhino species that were living in Europe and Asia toward the
end of the Pleistocene, but I must, in order to do so, say
a bit more about Bwana, the “tame” black rhino
we met in Chapter 3.
Despite all the difficulties which he has encountered in
the human world in which he was raised, Bwana’s worst
problems have been with other rhinos. One or two of his escapes
have taken him into the surrounding wilderness where he has
been injured by wild males. Bwana’s difficulties with
those males results from his ignorance of the basic rules
of rhino life. In the wild, only males who’ve established
dominance in a particular territory are allowed to spray clouds
of aerosol urine out behind them. (A rhino’s penis points
backward when it isn’t erect.) Those urine mists swirl
over the dominant male’s hindquarters and over the surrounding
vegetation to announce his status and, therefore, his sexual
eligibility. They convey a message which is, obviously, of
the highest importance in rhino society. Subordinate males
must urinate in a “non-aerosol” stream straight
down onto the ground to show that they have no intention of
usurping the sexual privileges of the “owner”
on whose property they’re living. If they don’t
urinate in this modest manner, the dominant animal will chase
them off, and kill them if they resist. Bwana has, however,
never learned these rules, so he’s always issuing unintended
affronts to the reserve’s dominant males.
Defecation is an even more complicated matter than urination
in rhino society. Rhino dung-heaps or middens are located,
Richard Estes’ Safari Companion tells us, “like
signposts along rhino paths and territorial boundaries.”
It seems that all or most of the rhinos in the area defecate
on them: the dominant bull, the subordinate males and visiting
cows. We don’t know whether subordinate bulls are merely
permitted to use these “latrines,” or
whether they’re actually obliged to do so.
Does use of the dung middens give them a kind of “residence
permit” in the dominant male’s territory? Does
it say to him, in effect, “this is to remind you that
I’m not a stranger here, and please note too, from the
hormonal signature of my excretions, that I’m not about
to challenge you”? I don’t know the answers to
these questions, and, for now, Bwana doesn’t either.
The people at Lapalala hope, however, that they will be able
to initiate him into the complexities of rhino life by placing
him in a remote part of the reserve with a few females and
younger males.
By far the most important thing that Bwana will have to learn
about dung midden etiquette is that the dominant male – and
only the dominant male – is allowed to perform
the Dung-Spreading Ceremony on them. He does this by dragging
one hind leg after the other sharply through the midden in
a gesture which is as formal and stylized as the battement
tendu in which a human ballet dancer draws his or her
toe across the stage floor. This battement tendu
is such a fundamental part of the rhino language of sex and
dominance, that males start practicing it as small calves.
The dominant bull raises his tail and defecates explosively
before or while he’s doing it, ejecting softball-sized
dung-spheres onto the midden in a compelling blend of drama,
beauty and scatological humor.
* * *
Like humans and chimpanzees, white and black rhinos share
a relatively recent common ancestor. Hominids and early white
rhinos split, in fact, from their more forest-adapted relatives
at roughly the same time in response to the appearance of
relatively grassy conditions in the late Miocene-early Pliocene
of 6-8 million years ago. It’s no exaggeration, therefore,
to say that white rhinos and hominids have been part of each
others’ lives right from their origins.
When members of Homo sapiens left Africa around
80,000 years ago, they encountered the still-existing Rhinoceros
unicornis, the one-horned Indian rhinoceros, as well
as two smaller species which are now, in the early years of
the twenty-first century, on the brink of extinction: Rhinoceros
sondaicus, the one-horned Javan rhino (a smaller variant
of the Indian rhino), and Dicerorhinus sumatrensis,
the small, hairy, two-horned Sumatran rhino.
They also encountered three rhino species which have become
extinct. The largest of these was a huge grass-eating rhino
called Elasmotherium sibiricum which lived in and
around what is now Russia. Even bigger than the white rhinoceros,
Elasmotherium carried a single, enormous horn whose
base extended from its nose to its forehead. The other species
that became extinct were two-horned rhinos. They were, respectively,
two woodland Eurasian species in the genus Stephanorhinus,
and a steppe species called Coelodonta antiquitatis.
Coelodonta, the so-called “woolly” rhino,
was a grass-eater similar to, but a little smaller than, the
white rhino. It occupied an enormous range stretching from
China to Spain. Between 17,000 and 13,000 years ago, a human
artist created this image of a Coelodonta rhino in
the so-called “Dead Man’s Gallery” of Lascaux
cave in southern France:
The late Björn Kurtén, one of the greatest authorities
on Pleistocene mammals, found the image puzzling: “What,”
he asks on page 125 of The Ice Age, “is the meaning
of... the six black dots?” In his The Cave of Lascaux – the
Final Photographs, Mario Ruspoli says they are “a
sign meaning the end.”
Coelodonta was a close relative of the two African
rhino species, and, from what I’ve seen of white rhinos
in the Kruger Park, (both in the flesh and in the excellent
video footage of their mating rituals that has been shot there)
it seems clear that the Lascaux rhino, with its right back
leg trailing in the battement tendu position and
its raised tail, is performing the Dung-Spreading Ceremony.
The six black dots are his dung balls. (The use of dung heaps
as territorial and sexual notifications runs deep in the rhino
family, and even extends to some of its surviving relatives
in the odd-toed or “perissodactyl” order, horses
and tapirs.) The fact that no-one else has, to my knowledge,
picked up on this interpretation of the rhino painting in
Lascaux doesn’t mean that I’m more perceptive
than observers like Björn Kurtén – it’s
an indication, rather, of how few present-day members of our
species have the opportunity of observing in the wild an animal
that was once such a familiar part of human life.
* * *
Fifteen thousand years ago, all humans grew up surrounded
by animals, birds and fishes. Children living in Europe at
this time could probably distinguish mouse, vole and shrew
species with an expertise which only a handful of specialist
zoologists possess today. Migrations of reindeer, salmon,
geese and swans would have marked the passage of their seasons.
Those children would have watched great auks extending their
wings for balance as they waddled across wave-washed rocks
on both the Atlantic and Mediterranean shores. (This image
doesn’t spring, by the way, from my imagination – great
auks are depicted in this position in paintings in the recently-discovered
Cosquer cave near Marseilles.) And no matter where in Europe
those ice-age children lived, the moaning of lions and the
whooping of hyenas would have been as familiar to them as
emergency-vehicle sirens and car alarms are to their present-day
counterparts. The presence of those lions and hyenas – and
that of leopards, brown bears and wolves – wouldn’t
have soured their day-to-day lives with fear, but it would
have kept them focused on the danger of wandering away from
their families after sundown.
Many of the mammals that lived in the Europe of fifteen thousand
years ago are still found there. Lynxes, wild cats, otters,
badgers and other mustelids, bats, rodents and shrews of various
kinds, foxes, wolves, bears, horses, cattle (called “aurochs”
in their wild form) ibex, chamois, reindeer, roe deer, fallow
deer, red deer, moose (called “elk” in Europe),
and bison (“wisent”) are all in this category.
Some of these survivors (like bison and wolves) hang on in
pitifully small numbers, while the cattle population has,
like that of humans, become hugely inflated.
Other mammals have disappeared from Europe, but are still
found elsewhere – lions, hyenas, leopards, saiga antelopes,
musk oxen and beavers are in this category. Still others have
disappeared completely: the giant deer Megaloceros,
a large vegetarian bear called Ursus spelaeus, the
Coelodonta rhino and the mammoth are in this category.
Mammoths are often lumped together with dinosaurs as “prehistoric”
animals, but dinosaurs became extinct sixty-five million years
ago, long before there were any big mammals at all, while
mammoths were thoroughly modern elephants which only evolved
about six to eight million years ago in response to the same
phenomenon which gave rise to humans and white rhinoceroses – the
Miocene drying-trend which stimulated the development and
spread of grasslands.
Grass is an excellent and abundant source of nutrition, but
it’s extremely tough and fibrous, and protected, in
addition, with abrasive silica. Mammals have evolved two main
strategies for processing it. One involves the development
a specialized digestive tract populated with bacteria that
have become increasingly good at breaking down its fibers
chemically. The other is the evolution of tough, specialized
teeth which can break those fibers down mechanically.
Mammoths, and their close relatives Indian and African elephants,
chose the latter path. The distant, insectivorous ancestors
of mammoths (and all other placental mammals) had 44 teeth.
Those ancestral eutherians arose late in the Mesozoic, or
age of the dinosaurs. Since the start of the Cenozoic or “age
of mammals,” there’s been a trend among almost
all their descendants toward reducing the ancestral number
of teeth. That trend is weakest, as one might expect, in a
mainly-insectivorous group of mammals which is still close
to the ancestral type. Solenodons, tenrecs, otter-shrews and
golden moles are still, therefore, equipped with 40 teeth,
while some hedgehog, mole and shrew species retain the full
ancestral complement of 44. Humans, apes, baboons and monkeys
– animals who’ve traveled an intermediate distance
from the ancestral point in this respect – have 32 teeth.
Rodents, whose dentition is more developed or “derived”
than that of primates, typically have 16. Mammoths and modern
elephants, which underwent what its perhaps the most extreme
degree of modification in this regard, went down to having
ten teeth at a time in their mouths.
Early mammals developed “specialized” teeth even
before they split into marsupial and placental branches. The
undifferentiated peg-like teeth of their reptilian ancestors
(designed for seizing and holding) evolved into incisors in
the front of the jaw, and into shearing-and-grinding teeth
in the cheeks. Only two seizing-and-holding teeth were retained
in each jaw – the “canines” that separate
the incisors from the cheek teeth.
At first, cheek teeth had three main cusps. A low triangle
formed by those cusps fit or “occluded” precisely
into the cusps and hollows of an oppositely-oriented triangle
on the opposing tooth. This “tribosphenic” arrangement
combined the shearing and grinding needed for an insectivorous
diet so advantageously that two branches of early mammals
may actually have evolved it independently. As mammals started
moving, however, into the herbivorous niches left vacant by
the extinction of the dinosaurs, grinding became more important
to many species than shearing. This prompted the emergence
of four-cusped or “quadrate” cheek-teeth. Human
cheek-teeth are quadrate – as are those of raccoons, bears,
pigs and ancestral elephants.
In certain species, some of the cusps on those quadrate teeth
have become linked by ridges or “lophs.” In primates,
for instance, each front cusp has become joined to the cusp
behind it. The lophs on our pre-molars have become so prominent
that it looks like those teeth have only two long cusps. They
are referred to, in fact, for this reason, as “bicuspids.”
Elephants also developed lophs between the cusps of their
quadrate cheek-teeth, but their lophs ran diagonally to ours.
Each cusp was, in other words, linked to the one next to it,
rather than the one in front of it. Evolution then started
adding new pairs of cusps – joined by new lophs – to
elephants’ originally “bilophid” molars.
“Trilophodon” elephants appeared, followed by
tetralophodons, and, later, by pentalophodons. This development
was to culminate, eventually, in the development in mammoths
of long, massive molars with up to sixteen lophs.
Back at the tetra- and pentalophodon stages, however, elephants’
molar cusps were still quite a bit higher than the lophs.
They rose above those connecting ridges in pairs of cones
which reminded early taxonomists of breasts (“mastodont”
means “breast tooth”). Despite the fact that new
pairs of cusps were being added to them, the basic design
of those “breast teeth,” remained the same as
that of our teeth: a dentine core, covered with an enamel
crown. Those enamel-covered cones worked fine on the leaf
and twig-dominated diet which elephants ate during the Miocene,
but something more resistant to abrasion was required to process
a steady diet of grass.
That “something more resistant” began to emerge
in the middle Pliocene as the lophs grew higher, fusing each
pair of “breasts” into a single ridge. The height
of those ridges then increased, and they got squashed together
as more and more of them were added to the tooth. The spaces
between them became filled, too, with cementum. The ridges
were then extended, along with their enamel covering – deeper
and deeper into the tooth. They became folded together so
tightly that they began to look like pleats, and the end result
was a tightly-packed series of buried enamel sheets which
filled the tooth and from top to bottom.
The complexity of the new molars made them so big that there
was only room for a few of them in the jaw. Gomphotheres had
already reduced their relatively simple “tetraloph”
cheek-teeth to twelve, and mastodonts went down to ten. The
Elephantidae (the suborder to which mammoths and the two surviving
elephant species belong) got down to eight cheek teeth–
a single “main grinder” doing the bulk of the
work in each quadrant of the mouth, together with a second
one “waiting in the wings” to replace it by moving
in from the back of the jaw.
The production of two sets of teeth – “milk”
or “deciduous” teeth followed by an adult or “permanent”
set – is such a common and widespread characteristic of
placental mammals that it’s thought to be an ancestral
condition. Like their distant relatives, manatees, elephants
have, however, departed from it – their cheek teeth are
replaced no less than five times. The first two teeth to appear
are premolars. The remaining three are molars.
All teeth other than molars and premolars have disappeared
in the Elephantidae, with the exception of two upper incisors
which are elongated into tusks which could be over 10 feet
long in African elephants, and up to sixteen feet long in
largest of the mammoth species, the Columbian. Those two incisors,
plus two cheek teeth at a time in each quadrant of its mouth,
give the Elephantidae its total of ten teeth at a time.
The Elephantidae aren’t the only mammalian group whose
molars are equipped with buried enamel plates – many rodents
also have teeth with plenty of lophs running across them,
and some (like South American capybaras and South African
whistling- and vlei rats) have squashed those lophs together
and buried them inside their molars to create the same “washboard”
or “loxodont” grinding surfaces that we see on
elephants’ molars.
Even though savanna elephants eat plenty of bark, branches
and leaves, the grass which stimulated the development of
their huge and complex molars is still a very important item
on their menu. The Asian elephant and the cyclotis
subspecies of the African elephant have returned to a forest-dwelling
existence, but, regardless where they live, all existing elephants
still possess the specialized tooth structure that made it
possible for their suborder to take possession of the grasslands – the
velds, steppes, pampas and prairies of the Pleistocene.
* * *
The living members of the Elephantidae give us a very good
idea of how mammoths must have conducted their social lives
and raised their offspring. The two still-living elephant
species show us, too, how an animal the shape and size of
a mammoth or a mastodont must have moved – what
its living appearance might have been.
Hoping to get a similarly lively clue to the movement and
appearance of the giant Megaloceros deer we spoke
about a few paragraphs back, I followed a group of eland that
was moving along the opposite bank of a ravine in Suikerbosrand
nature reserve near Johannesburg. Moose, which top out at
around 1,750 pounds, are currently the biggest deer in the
world, but the eland, a giant antelope, can weigh up to 2,000
pounds. I wasn’t as impressed as I was expecting to
be by the size of those eland, until they were joined by what
looked like a herd of miniature zebra. Only then did things
shift into perspective: the “miniature” zebras
were, of course, normal-sized animals, and the eland were
suddenly revealed for the unlikely giants that they really
are. A second group of eland, a bachelor herd which I saw
at much closer range on Suikerbos’ Bokmakierie hiking
trail, gave me a better look at the thick legs and massive
bodies of this extraordinary species. The dewlaps hanging
from the bulls’ bulky necks reminded me of similarly-equipped
domestic cattle. The thick tufts of hair between their eyes
gave their foreheads a bulbous look in profile, and I could
hear, for the first time, the clicking noise the feet or legs
of these giants make when they walk.
Imposing as they are, eland still weigh only about seventy
percent as much as Megaloceros did. Seven feet tall
at the shoulder, the giant deer carried antlers 12 feet wide,
and weighed somewhere between 2,500 and 3,000 pounds – as
much as a big giraffe. Because the top speed of big animals
tends to be absolutely faster, but relatively slower than
that of small ones, (the naval architect William Froude described
the same phenomenon in relation to ships and, indeed, waves)
a running Megaloceros must have had the intriguing slow-motion
quality we see in a galloping giraffe.
* * *
Europe’s animal inhabitants divided that continent
among them on a “timeshare” basis. During the
cold parts of the glacial cycles, the continent was occupied,
for the most part, by cold-adapted animals like mammoths,
woolly rhinos, and reindeer. During the warm “interglacials”
we’ll talk about in Part 3, this “cold-weather
fauna” retreated to the northern reaches of Eurasia,
while present-day “middle-Europe” passed into
the possession of warmth-loving animals like the straight-tusked
elephant Paleoloxodon, the woodland rhinos Stephanorhinus
hemitoechus and S. kirchbergensis, and the hippopotamus
species that still lives in Africa.
Throughout the last million years, these “warmth-loving”
animals had been driven southward many times by the glacial
episodes that have been occurring every 100,000 years on average.
These glaciations either pushed them right out of Europe to
find refuge in Turkey or the Middle East, or restricted them
to Europe’s Iberian, Italian and Balkan peninsulas.
After each cold episode, they would return to middle-Europe,
as warm, moist interglacial conditions spread broad-leafed
forests back into that region.
The warmth-adapted animals did not, however, return, after
the last glaciation, because all of them, (except Hippopotamus
amphibius which survived in Africa) had become extinct
by about 25,000 years ago. Paleoloxodon was already
gone by about 50,000 years ago. It’s highly unlikely
that the latest glaciation killed these animals off. They
had, as we’ve just seen, successfully survived all the
previous Pleistocene glaciations – some of which were
more severe than the last one. The only unique thing about
the last glaciation, was the fact that Homo sapiens
took possession of Eurasia in general, and their relatively
warm glacial “refugia” in particular, while it
was in progress.
* * *
Europe’s extinct megafauna included a member of the
human family. That species had evolved from beings identical
with, or very similar to, Homo erectus. These erectus-like
beings, which are also referred to as H. heidelbergensis,
probably continued to exchange genes with their Asian and/or
African relatives from time to time. By the height of the
great Riss glaciation some 150,000 years ago, they had, however,
evolved a body-shape distinctive enough to differentiate them
from their immediate ancestors. In 1869 William King named
that new species H. neanderthalensis.
Heidelbergensis-neanderthalensis had, as we’ll
see in Chapter 12, been expert and regular hunters of big
game right from the time they entered Europe. They also knew
how to make fire, and used their stone tools to fashion sophisticated
spears and other implements out of wood. Between 350,000 and
410,000 years ago, at Bilzingsleben in east-central Germany,
a heidelbergensis individual incised a pleasing,
abstract pattern onto a bone artifact, whose discovery made
the chief researcher at that location, Dietrich Mania, suspect
what he calls “the development of a capacity for abstract
thought, and the presence of language.” (“...die
Herausbildung der Fähigkeit zum abstrakten denken, und
das Vorhandensein einer Sprache.”)
The discovery of the incised patterns constitutes a particularly
significant enlargement of our understanding of heidelbergensis,
because the production of decoration and art is regarded – even
more jealously than some people regard the use of bone tools
and the ability to hunt big game – as an exclusive province
of Homo sapiens. Although there is no evidence at
all that the heidelbergensis -neanderthalensis line
produced representational art, it does not seem to have been
indifferent to decoration and adornment: ochre and manganese
oxide pigments have been found at several sites associated
with heidelbergensis, while ochre and ochre-grinding paraphernalia
have been found at sites occupied by Neanderthals.
Our species of human, Homo sapiens, first entered
Europe between 45,000 and 50,000 thousand years ago, probably
via what is now Bulgaria. By 38,000 years ago, sapiens had
reached present-day Portugal. A second wave of sapiens, equipped
with the so-called “Gravettian” complex of technologies,
entered Europe some 30,000 years ago from what is now Russia.
By approximately 28,000 years ago, neanderthalensis
became extinct. Mammoths and woolly rhinos disappeared from
Western Europe around 12,000 years ago, the former persisting
in Siberia nearly two thousand years longer.
Chapter 5
- Lost
Serengetis 2: Australia and North America
