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Jonathan Kingdon:
Lowly Origin:
where,
when, and why our ancestors first stood up
(Princeton University
Press: 2003)
“Speciation, especially
multiple speciation, has to take place in geographic or
ecological compartments.... All animals have finite distributions
that are loaded with many detailed implications for their
ecological adaptations, evolutionary origins, and ability
to spread or disperse. Both contemporary and ancient Africa
can be understood as a pattern of ecological islands....
Discussion of the geography of human evolution has often
been so threadbare, abstract, and generalized that our
many and different ancestors have no perceptible existence
in time and space. There needs to be a fuller acknowledgement
and awareness that our forebears were embedded in the
same ecological matrices that other animals are and have
been, all with specific and finite distributions.”
(Kingdon, p.10)
This, to put it bluntly, has been the major weakness in
previous hypotheses regarding the early evolution of hominids.
“And then they went out onto the savanna & stood
up” is, implicitly or explicitly, the basic story
in dozens of books, and literally thousands of scientific
papers. The one thing that virtually no-one seemed really
prepared to do was to treat our common ancestor as just
another big mammal - which, by definition, it was - and
apply the full range of biogeographical information to
the problem of its origin. Similarly, standing upright
requires a whole host of anatomical shifts, which must
have occurred as an ongoing adaptive process that was
viable throughout...otherwise, no descendants.
Putting these two rich sources of information together
is the key innovation in Jonathan Kingdon’s Lowly
Origin, a work which has clearly set a new benchmark
for approaches to human evolution. Quite simply, it outclasses
all previous attempts at answering this question, making
them appear simplistic & naive in their failure to
address the biogeographical context, or the genuine complexity
of the anatomical rearrangement that was required.
Kingdon is amply qualified for this work. A leading expert
of African mammalian ecology and anatomy, he is also a
very good writer & a subtle thinker who is at ease
with multicausal explanations - which he constantly ends
up with, despite his efforts to distinguish between diffferent
adaptations, in an attempt to make the processes involved
clearer to the reader:
“One difference
between this account and many that have preceded it is
that I try not to amalgamate adaptions.... A second peculiarity
of my own understanding of human evolution can be contrasted
with the pictures that are painted in innumerable books,
articles, and dioramas representing hominins and other
extinct mammals in picturesque ‘National Park-like’
fire-climax savannas...[which] became common only 1 to
2 mya [million years ago].... Another primary difference
between my approach and those of my predecessors is that
I envisage standing as a relatively inefficient response
to an exceptionally benign but very localized environment,
This is the exact converse of previous explanations, which
attempt to understand bipedalism in terms of increased
efficiency under very widespread ‘savanna’
conditions that were more difficult and trying than those
in the forests or woodlands that preceded this supposed
‘ordeal’.... [As well] there have been various
suggestions for the isolation of vaguely eastern or southern
ape populations, but none has identified a habitat both
ecologically distinct enough to elicit an entirely new
form of locomation, nor geographically separate enough
to impose the necessary isolation. My own outline addresses
both shortcomings.”
(Kingdon, pp.14-18)
As is made evident here, not only is the typical “savanna”
model likely to be incorrect, it is literally impossible
- as such landscapes only became common enough to be a
major evolutionary factor much more recently...and, quite
possibly emerged - as we shall see - only in the wake
of “firestick farming”.
“For the most
part, I have used the often random and accidental provenances
of fossils as mere guides to the larger ecological and
geographic contexts for human evolution, seeking clues
in those details of African biogeography and ecology that
we can still retrieve and reconstruct today. I have also
sought to put the likely anatomical and behavioral responses
of early hominins to a succession of environmental challenges
into a sequential and spacial order that is consistent
with the fossil record.... The first tie-up between time,
place, ecology, and behavior is located on the east African
coast, the second and third involve movement into the
interior (each involving subtly different but highly significant
divergences). The hominin trail leads on into the Highveldt
and other interior uplands and thence, very much later,
to the Atlas Mountains (or Arabia).... Finally, as a specialist
in the evolution of mammals, the perspective that I have
sustained the longest (and reinforced most decisively
in this book) is that of the emergence of humans as the
evolution of yet another mammal - a very peculiar and
special one, true, but in essence just one more African
mammal.”
(Kingdon, pp.5-6)
“As for mechanism,
I have not sought global drought crises, nor even fewer
trees. Rather, I see an ape population adapting to a different,
more deciduous kind of forest and...different, perhaps
richer, menu. I have postulated a switch to more terrestrial
feeding, but instead of actively pursuing fleet prey,
east coast ground apes would have found a rich supplement
of small, sessile animals and plant matter from the forest
floor to augment crops of fruit; the latter being predictably
less diverse and growing nearer the ground than in the
high forests further west.... The argument hinges on changes
in the spine, pelvis, and head-neck junction (perhaps
also the heel) being necessary precursors to standing
and balancing on two legs.... In common with my colleague,
Clifford Jolly, I hold that it was foraging, mainly on
the ground, in a squatting position that demanded these
necessary modifications.”
(Kingdon, p.19)
By this stage, readers should be under no illusions. This
is not the usual
tale of human evolution they are used to. In parallel
with Merlin Donald’s Origins
of the Modern Mind (1991) - which redefines cognitive
evolution - I would argue that Kingdon’s work ought
to decisively reorient human evolutionary approaches towards
much more detailed and coherent models, which fully think
through the implications of our ancestors as real, functioning
animals, who evolved in specific ecosystems that are hardly
likely to be well represented in the modern world.
Kingdon also wants to reconceive the debate surrounding
the freeing-up of our hands through upright posture, by
exploring the long different nature of fore- and rear-limbs...something
that was obscured when vertebrates first took to the land:
“From the very
beginning, tail-end limbs have had quite different origins
and functions from forelimbs. The latter first evolved
in close association with the head - indeed, so close
that pectoral fins were actually tethered to the head
of early fishes.... Likewise, the anterior sources of
forelimb nerve networks testify to the very separate limb
origins. Subordinate to the brain, forelimbs still serve
many vertebrates to alter and adjust their sense-driven
decisions about the direction and pace of forward movements...
Even in species with well-matched limbs and great similarities
in hand and foot structure, a clear separation of functions
between fore- and hindlimbs is evident. All four extremities
function as clamps, but there is a premium on both strength
and ankle flexibility in the rear clamps whereas ‘exploratory
skill’ and deft, multiple use distinguish the fore
clamps.”
(Kingdon, pp.26, 50)
“A reasonable
working hypothesis is that primates arose from the already
aboreal and nocturnal [mammalian] stock as a peculiar
lineage of tiny visually oriented insect eaters, foraging
at night, singly, and probably rather slowly, through
the fine foliage of tropical rainforests.... Realignment
of the eyes seems to have been an intrinsic part of the
muzzle’s shrinkage, an expansion of the brain, and
a downward bending of the skull and shortening of the
face.... In an evolutionary perspective, [primate visual]
improvements essentially redifferentiated and regained
properties of vision that were never lost by birds, reptiles,
even some fish!”
(Kingdon, pp.45-6)
“I think that
what took place was less a case of bipedalism initiating
new behaviors, than the removal of frustrating constraints
on many existing talents. These could be said to have
been warped or at least hampered by the anatomy of a weight-bearing
wrist and hand. Before becoming bipedal, the potential
for more effective manual manipulation of foods or fellows
must have been curtailed by the persistent intrusion of
weight-bearing duties.”
(Kingdon, p.126)
By setting the scene in this way, Kingdon frames the problem
afresh. Rather than intensified selection pressures
on locomotion leading the way, these were relaxed whilst
the underused potential of the hands were mobilized afresh.
Given the sheer complexity of the redesign of the spine,
pelvis and shoulder involved, this appears a considerably
less “miraculous” adaptive process, and one
well-fitted to the now almost lost ecosystem Kingdon identifies
as our ancestral home...
“It was squat-feeding,
not bipedalism, that induced changes in the upper body,
backbone, and pelvis of ground apes. The special legacy
of squat-feeding was to disengage a heavy, cantilevered
upper body from an equally oblique pelvis, and to rebalance
a more lightly built head and thorax vertically over a
compact basinlike pelvis. That these changes could improve
balance on two legs would have been an almost accidental
bonus, an anatomical by-product.”
(Kingdon, p. 153)
“If the bifurcation
within a common chimp-hominin ancestral population took
place in eastern Africa, it is essential to explore how
habitats in central Africa might have related to those
in the east at the same time. In the Horn, recurrent cycles
of drier climate have repeatedly interpolated an ecological
barrier between east-central Africa and the humid coast....
[It was] climatic change operating through this Somali
arid zone, rather than the Rift valley, that premitted
protohominins to become a new, highly distinctive and
aberrant primate.”
(Kingdon, pp. 106-7)
“Kiwengoma, in
the Matumbi Hills on southern Tanzania’s Indian
Ocean littoral, is a last vestige of the forests that
one capped hilltops and filled valleys all along the east
African coastline. The distinctness of this habitat has
only begun to be documented and appreciated in very recent
times...as a ‘Center of Endemism’ quite distinct
from the main forests of central and western Africa.
That distinctness lies not only in the different climates,
soils, and species. For the animals and plants that live
there, the narrowness of the coastal strip implies both
diminished population sizes and much greater susceptibility
to fragmentation.... [As well] the distribution of many
plants and animals testifies to the east coast being the
core area for forest and thicket regimes that spread far
inland, mostly along broad valley systems and up the moister
slopes of inland hills and mountains.”
(Kingdon, pp.116-18)
And the innovative ideas (and the evidence to support
them) just keep coming. Kingdon makes no bones about the
fact that the fossil record is very likely to provide
only a sample of the plethora of early hominin species,
based upon the species radiation patterns we observe in
other large mammals. He also makes a strong case connecting
this with the environmental context he describes:
“If my suggestion
is correct that the hominin radiation began with a coastal
stock of ground-living apes...a discontinuous distribution
over some 4000 km may well have given rise to genetic
differences [even] in a purely coastal species. As soon
as that parental population began to expand inland from
its coastal base, the possibilities for genetic differentiation
would have multiplied.”
(Kingdon, pp. 157-8)
“’Last ape’
to ‘first hominin’ is only a conceptual transition
(and a largely semantic one at that), but there could
be a convenient geographic dimension whereby the former
remained ‘squatters’ that lived in moist,
warm, nutritionally rich coastal lowlands, whereas the
latter were (probably rather slow) ‘walkers’
that took to a somewhat more exposed way of life beside
rivers in the drier, cooler, and higher uplands.... During
later periods of global cooling, these upland populations
may well have been forced to descend from the heights,
but an acquired tolerance of cold and drought should have
put south African and Ethiopian populations or groups
at an advantage.”
(Kingdon, p.164-5)
By making substantial use of the observational work of
primatologists, Kingdon also develops a useful - and likely
- approach which sets the stage for proto-culture firmly
within the capacities of a band of upright apes. And,
at every stage of his argument, the clear evidence from
a variety of sources - supporting overdetermined shifts
in highly likely directions - sets, yet again, a new standard
for such work.
“In spite of a
menu of behavioral options and some potential for learning
refinements, the displays of most species (including ‘bluffing’
or ‘mobbing’ routines) are stereotyped and
genetically fixed. Apes, instead, can elaborate or adapt
both individual and group actions...[which] can justifiably
be described as cultural, learned behaviors.... Scavenging,
in the popular sense, could have been the late manifestation
(after Homo had
become a full member of the open country fauna) of a much
larger trait in our lineage. This trait has less to do
with a taste for meat from dead animals than with particularly
acute sensitivities to other animals as guides to hidden
or potential food sources. This sensitivity could have
been to the animals themselves, as prey, but progressively
tended, more and more, toward appropriating some or all
of the other animals’ own subsistence...displacing
or discouraging any other species from food sources they
might otherwise be able to use. Scavenging, as commonly
understood, is typical of open country species. I contend
that its beginnings, in forest-based members of the Homo
lineage, could have provided a major mechanism not only
for expansion out tof the forest, but for a diversification
of diet that led away from species-specific diets and,
even more significantly, a rapid elaboration of flexible
technologies for obtaining food.... Combining the predator’s
alertness to other species’ behavior with a periodic
intolerance of actual and potential competitors (as food
choices increased), this unprecedented foraging technique
might better be termed ‘niche-stealing’ than
scavenging.”
(Kingdon, pp.214-19)
If Jonathan Kingdon is right, and I strongly suspect that
he is, our complete picture of early human evolution will
have to be totally redrawn. Even so, his book does not
address all the issues - concentrating on material rather
than psychological processes - making it particularly
strong on the period up to the emergence of Homo
ergaster/erectus. As I noted early in this review,
perhaps the best counterpart to Lowly Origin is Merlin
Donald’s Origins
of the Modern Mind. For there is very little overlap
between the two, whilst they complement each other marvellously.
However, despite the major contrast in focus, and their
almost entirely different disciplinary backgrounds, there
is one main feature which unites the two, and which is
sadly lacking in most treatments of human evolution. This
is the insistence, by both authors, that we need to develop
models that envisage our ancestors as fully functional
& integrated beings, not just some half-way house
between apes and humans. And, although this principle
attracts a great deal of lip service, it is rarely treated
as the difficult task that it is...particularly at the
level of Homo ergaster/erectus.
Equidistant from us and our closest cousins - the two
chimpanzees - erects colonized Eurasia, mastered fire,
and developed the first highly-worked technology. Yet
their anatomy strongly suggests that they could not speak,
despite their substantially enlarged brains, and the evidence
is clear that their culture was basically static compared
to ours.
“It will be clear
enough now that my piecemeal approach to hominin evolution
includes the possibility of a ‘slow start’
for the Homo
lineage (relative to the Praeanthropus /Paranthropus
group, which I regard as belonging to a more precocious
sibling lineage).... Further, I am suggesting that the
most radical transformation could be be ascribed to an
outlying population suffering a relatively late but lengthy
isolation in a distant, dry, open habitat.... The association
of ergaster with
fire is of exceptional interest, because scholars have
long ascribed the reduced size of human teeth to cooking
and food processing, and it is the small cheek teeth that
most immediately distinguish H. ergaster ....
[And] if the Atlas region was this species’ region
of origin, it can be remarked that (in common with some
other Mediterranean environments) fires triggered by lightning
are an annual hazard....[Moreover] the link between H.
ergaster may be much
more extensive than has been appreciated. What are commonly
called ‘secondary grasslands’ are in fact
made by two principal agencies: one is grazing, browsing,
and trampling by large or numerous herbivores; the other
is fire.... Thus, the spread of fire-climax grasslands
may not have preceded the burgeoning population of highly
successful hominins, but could have followed hard on their
pyromaniac heels.”
(Kingdon, pp.271-7)
Lowly Origin
reconfigures the entire frame for debating human origins,
by immeasurably enriching our understanding of the context
in which it occurred - and the exact nature of the anatomical
changes involved. There is simply no substitute for this
book...and until other researchers come near to matching
its sophisticated grasp of these issues, it will be difficult
to say whether other evolutionary scenarios can match
it. Kingdon’s book is that most rare thing - genuinely
original, and the best in its class. Read it...
“As yet, there
is no direct evidence for the detailed population structure
or demography within groups of prehistoric foraging humans....
Up to a third could have been less than fully adult (in
spite of high
mortality or low birth rates), which would have given
this class some sort of proportional role in determining
the day-to-day functioning of the group. It would be wrong
to see nonadults as mere hangers-on...they must have been
active participants in the overall foraging.... A high
proportion of nonadults defines what might have been special
about human evolution. It also hints at where the power
of natural selection is targeted and is most influential.
In thinking that selection favors smart kids and loving
mothers, I depart from most other students of human evolution,
who have concentrated on sexual selection and mate choice
in their search for primary selection mechanisms.”
(Kingdon, p.294)
John Henry Calvinist
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