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Human Activities and Megafaunal Extinctions: Man's modification of the environment may
have caused the demise of some large Pleistocene mammals
Author(s): Grover S. Krantz
Source: American Scientist, Vol. 58, No. 2 (March-April 1970), pp. 164-170
Published by: Sigma Xi, The Scientific Research Honor Society
Stable URL: https://www.jstor.org/stable/27828993
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 Grover S. Krantz
One of the most vexing and most dis
cussed problems that researchers in
both paleontology and archeology are
concerned with is that of the extinction
of large animals in the Late Pleistocene.
For all the efforts to clarify the cause
or causes, there remain two sharply
opposed schools of thought. Some au
thorities hold that the hunting activity
of prehistoric man directly eliminated
many species; others believe their
disappearance is primarily a result
of abrupt climatic changes. Both views
are well represented in the recent
volume edited by Martin and Wright
on Pleistocene Extinctions (1967). The
present paper offers a new interpreta
tion of the data, which might have had
a place in that book had the study
been undertaken a few years earlier.
Man may well have caused many of
these extinctions, I believe, but not by
the direct means of physical elimina
tion through excessive pr?dation. I
shall describe below three possible
indirect means: (1) competition for a
particular food supply, (2) causing
one herbivore to exterminate other
herbivores, and (3) agricultural prac
tices altering ecological relationships.
The first and third methods have been
touched upon by others, while the
second, to the best of my knowledge, is
set forth here for the first time.
First let us examine the evidence that
prehistoric man may have hunted out
After getting his M.A. in anthropology at the
University of California, Berkeley, Grover
Krantz worked for six years at the Lowie
Museum of Anthropology in Berkeley as
preparator and technician. He has taught
physical anthropology courses, and completed the
requirements for the Ph.D., at the University of
Minnesota. His teaching and research center on
the origin and evolution of man. Address:
Anthropology Department, Washington State
University, Pullman, Wash. 99163
164 American Scientist, Volume 58
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Human Activities and Megafaunal
Extinctions
Man s modification of the environment may have caused the
demise of some large Pleistocene mammals
these animals. The most convincing
argument is the coincidence in time of
the arrival of man with these extinc
tions, or, as in Africa and South Asia,
the supposed first development of
big game hunting. Martin (1967: p.
Ill) gives the approximate dates for
these events in each area as follows :
Africa, South
Asia 40,000 to 50,000 years b.p.
Australia 13,000 years b.p.
Northern
Eurasia 11,000 to 13,000 b.p.
North America 11,000 b.p.
South America 10,000 b.p.
West Indies "Mid-Postglacial"
New Zealand 900 b.p.
Madagascar 800 b.p.
Another argument advanced against
any nonhuman or natural cause is the
lack of replacment of the extinct
forms by ecological equivalents.
Martin (1958) and Jelinek (1967) see
this as indicating human agency,
which alone could eliminate a species
leaving its ecological niche open.
However, Guilday (1967) shows how
the same phenomenon can occur under
the stress of temporary climate change,
as in the altithermal.
The last major argument is not so
much support for man's being the
exterminating agency as denial of the
most obvious alternative explanation?
climatic change. The nearly perfect
coincidence of so many mammal ex
tinctions in Northern Eurasia, North
and South America, and Australia
with the final withdrawl of the Wiscon
sin-Wurm ice sheet has led many
authorities to conclude that abrupt
changes in climate were the major
causal factor (Guilday 1967; Slaughter
1967; Hester 1967; and Kowalski
1967). This conclusion has been
challenged by those who point out that
no comparable extinctions occurred at
the ends of any of ihe previous glacial
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advances (Edwards 1967: p. 144;
Jelinek 1967: p. 193; and Martin
1967: p. 81).
Other, less often stressed, arguments
include the comparative absence of
extinctions among smaller mammals
in the Late Pleistocene as well as no
notable loss of plant species (Martin
1967: p. 78 and Leopold 1967). Had
climatic change been the predom
inant factor, it is argued, plants and
small mammals ought to have suffered
to some degree as well.
The above arguments could be ex
tended and documented at length, but
the same question always arises: some
new factor must have been present at
each time and place of massive extinc
tions?and the only new factor that
appears to be consistently present was
man. The conclusion seems inevitable
that prehistoric man must have hunted
and killed these animals on such a
scale that they were totally eliminated.
However reasonable this conclusion
may appear in the light of one class of
evidence, it fails utterly when ques
tioned from another angle: just how
was it possible for prehistoric hunters
to accomplish this feat? Even Martin
acknowledges the problem, stating
(1967: p. 115), "We must beg the
question of just how and why pre
historic man obliterated his prey," and
further, "The thought that prehistoric
hunters ten to fifteen thousand years
ago exterminated far more large
animals than has modern man with
modern weapons and advanced tech
nology is certainly provocative and
perhaps even deeply disturbing." It is
more than disturbing, it is impossible.
In order to clarify this problem some
what one must consider the presumed
causes of extinctions, in particular
the predator-prey relationship.
When a hunting species specializes in and in some respects even more surely. tinct animals from kill sites has been
taking a particular species as its food Man's hunting practices are entirely noted as a serious objection to the
source, the population of the hunter learned by each generation from the idea of man's role in their demise
is directly dependent upon that of its preceding one. Any break in this cul (Jelinek 1967: p. 198; and Guilday
prey. Only a certain number of in tural continuity would leave man with 1967: p. 137).
dividuals of a prey species can be out any instinctive behavior to depend At this point we seem to be faced with
"harvested" without altering its pop upon. Also, the special activities and two "facts" that are reasonably well
ulation level. This "harvest," in turn, knowledge required for hunting each
proven. First, the advent of big-game
will support only a certain population species of game will differ somewhat hunting man alone correlates with the
of the hunting species. If the prey from those needed for every other. times of massive extinctions. Second,
declines in numbers, so will the hunt Should one game animal become
hunting man could not and did not
ers, who can exist only in proportion scarce and thus more rarely en exterminate these animals. These two
to their food supply. Normally a stable countered, human hunters, in concen "facts" appear to be contradictory.
balance is maintained easily and auto trating more on other species (or Other researchers have proceeded on
matically. Some arctic species under upon vegetable gathering), will quickly the assumption that if one of these
go drastic fluctuations in population, lose the learned skills pertinent to "facts" is true the other must some
with cycles of several years' duration. taking the scarce game. Unlike in how be explained away or ignored.
Here too, the numbers of the hunting herent behavior programming, learned
species rise and fall correspondingly, skills can disappear in a generation, I shall attempt to show here that there
always in response to the availability and even seriously deteriorate in just is not necessarily any contradiction
between the two "facts." At the root
of the prey. a few years if not reinforced by
practice. of the matter is the assumption made
This relationship still holds true even by most authorities that man could
if the hunter is dependent on a partic In addition to the above points, non have exterminated a species only by
literate hunters are also known for directly destroying virtually all of its
ular prey species only during one brief
season of the year. However great a deliberate game conservation beyond individuals. There are other methods.
variety of food they may consume in the automatic system which applies to
Modern studies of ecology indicate
all other seasons, the hunter's popula other carnivores (see Heizer 1955 for that the major, if not the sole, cause of
tion will be limited to those who can be numerous examples). All primitive extinction is the removal of some essen
fed in the season of scarcest game. hunters are quite aware of the im
tial aspect of the environment, and
portance of females and young for
Clearly no extinctions will occur under future game supplies and will often not pr?dation. This sort of thing has
these circumstances. The scarcer the been suggested but little explained by
treat them accordingly?something
prey, the scarcer the hunter, and pr? no natural carnivore will do. some who refer simply to the "in
dation pressure eases off automatically. direct" influence of man (Simpson
The seeming wastefulness of cliff 1965: p. 228) or to the upset of a
If the hunter species is less particular drives may not be as significant as nicely balanced state of nature (Romer
and takes its food from a number of may appear. In the first place, such 1933 ; and Colbert 1942). The problem
game sources in all seasons, then a drives are not known to have been at hand is thus changed to answering
used earlier than 9,000 years ago
slightly different relationship follows. the question: What actions of man
Should one of the prey species be (Hester 1967: p. 181), somewhat too could have so altered the "balance of
hunted until its numbers are signifi late to be involved in most of the nature" as to deprive numerous species
cantly reduced, individuals of this major extinctions. Also, such stam of some ecological necessity?
species will be less often encountered. pedes are not age-selective in their
If the hunter is to maintain his popula destruction; they take young and old Direct Competition
tion independent of this reduction in alike, a point which will be seen later One distinct category of extinctions
one kind of prey, he must concentrate to have some significance. was, in a sense, caused by man: the
correspondingly more on other species. Direct evidence also exists that, in extermination of certain large carniv
As the hunter takes what are available orous mammals. Several writers have
North America at least, early hunting
of his several game species, those few man did not significantly prey upon pointed out how difficult it would have
est in number will be the least often been for early man to seek out and
those animals he supposedly exter
met and least often taken, thus tend
minated. Many archeological kill sites
destroy such dangerous animals as
ing automatically to be preserved which are now well documented in extinct lions, cave hyenas, and saber
from extinction. toothed cats (Eiseley 1943a ; and Butzer
clude the remains of animals killed by
Paleo-Indians in the Late Pleistocene. 1964: p. 400). It is all the more un
Vulnerability to pr?dation is thus not likely that man could have directly
an inevitable or even a likely cause of These data, summarized by Hester eliminated any such carnivores in
a species' demise. Extinction threatens (1967: p. 180), show that only two view of the predator-prey relation
only if the numbers are so reduced genera were hunted in significant
numbers, mammoth and bison, one ships noted above, considering the
that mating encounters utilize but a carnivores as prey and man as the
extinct and the other not. In 25 of the
small portion of their reproductive 34 documented sites bison was the predator.
potential. It is only with the advent of
"civilized man" that this becomes a exclusive or predominant game found, At least two writers saw why these
mammoth ran a poor second as the carnivores disappeared, though with
reasonable possibility.
sole or major game in 7, and at only 2 out suggesting the hand of man in the
These predator-prey relationships hold were various other animals empha process. Guilday (1967: p. 122) and
when primitive man is the predator, sized. The virtual absence of most ex Vereshchagin (1967 : p. 392) noted that

1970 March-April 165

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with the loss of their large herbivorous kill" is perhaps exaggerated. Mason occurrence in natural deposits hardly
prey the carnivores similarly vanished. (1962: p. 243) lists only 15 genera support the contention that man killed
Edwards (1967: p. 146) quite simply with a total of 20 species, and Newell them off. In order for man, or any
explains that as man can be a more (1963: p. 48) gives 12 genera and a other predator, to destroy a species
efficient predator, the others are total of only 16 species. Removing he must dispatch the vast majority of
eliminated by direct competition (see carnivores from these lists will reduce the individuals himself, and almost
also Martin and Guilday 1967: p. Martin's list by 4 genera, Mason's by all of these at an early age prior to
33-34 on Smilodon). This becomes a 2, and Newell's by 1. their reproduction. Clearly this did
case of ecological replacement: man The accuracy of these figures is per not happen, as the species in question
need not kill a single animal to elim haps worthy of a short digression. are well represented by adult animals
inate the species; he needs merely to They are based essentially on a com in deposits unrelated to any human
deprive them of their normal food parison of living fauna with the fossil activity.
supply by getting it first. This would fauna of the Upper Pleistocene. Earlier
If one accepts the archeological and
be especially effective at times of extinctions are based on comparisons
seasonal crisis, like northern winters, paleontological evidence at face value,
of one fossil fauna with another fossil
when food resources are reduced to an interesting conclusion seems to be
fauna. Since living and fossil animals indicated: that heavy hunting by
a minimum and competition is most have generally been classified accord
intense. early man preserved at least one type
ing to different procedures and by while unhunted species became ex
Probably most large carnivore ex different people, there is good reason tinct. This is just the opposite of
tinctions since the Middle Pleistocene to question any comparison of one over kill as it has been presented. It
were caused by the direct competition
with the other. Kurt?n has noted appears that while man preyed heavily
of man's increasing hunting abilities. (1968: p. 40) that the tendency now on the bison it increased in numbers
The early disappearance of saber in paleontology is to "recognize and range. To suggest that hunting by
tooths from most of the Old World was broad, inclusive species with a span man actually caused such an expansion
not from actual extinction of their of morphological variation similar to of the bison may seem odd, but this
prey species, some of which still that found in related species today," can be demonstrated to be a likely
exist, but rather from a scarcity of but that this has not been the practice possibility. The demonstration in
these individual animals (young, lame, in the past. There were probably far volves principally a comparison of
fewer extinctions at the end of the
ill, and aged) which they normally human pr?dation practices with those
preyed upon. These were being in Pleistocene than is usually supposed of other carnivores. Demographic data
creasingly taken by early human because there were actually not so on age distributions within a prey
hunters. The late occurrence of Macha many species which could become population often show sharp differ
extinct.
irodus in the latest glacial stage in ences depending on the type of pred
Britain (Flint 1957: p. 456) indicates It is likely that the American bison ator that is chiefly concerned.
their survival only in the coldest was a single species at any one time,
northern climates, which man could No satisfactory detailed figures seem
and that changes observed are a com to be available for bison, but the fate
not inhabit up to that time. Hunting bination of evolutionary change and of other animals in similar circum
man and sabertooths competed so movements of subspecies in response stances illustrates the point quite well.
closely that when the former was to climate shifts (Eiseley 1943b; and
present these great cats were not. Large natural carnivores prey mainly
Cornwall 1968 : p. 184). In my opinion on two age categories of their game,
Similarly, Smilodon vanished as soon Canis dims is probably not specifically
as big game hunting began in the with clear seasonal emphases. In
distinct from the living timber wolf ex
the warmer half of the year, beginning
New World, and shortly before its cept possibly as an ancestral species.
prey species died out. with foaling or calving, the young
The number of species of elephants re herbivores are the major part of their
ported in the literature is probably also diet. In the colder months it is mainly
Indirect Causes too many.
the oldest individuals, especially dis
So far I have discussed the role human
In spite of all such reservations it is eased and injured ones, that are taken.
agency may have played in extinctions still clear that a number of distinct At all times it is the prey that is
by somehow removing the food of a types of large herbivores did become easiest to catch that is eaten.
species. The next method to be de extinct within a short period of time.
scribed is even less direct: man may This kind of seasonal variation in pr?
These include mammoth, mastodon,
have caused certain herbivores to dation habits can be shown on a graph
horse, camel, several ground sloths, of age distributions of the prey
exterminate other species with similar at least one pronghorn, and possibly
ecological requirements. In this sec some cattle. population. A "survivorship curve"
tion I will concentrate mainly on the would show how many individuals re
well documented extinctions which The archeological record in North main at the end of each year from an
occurred in North America at the end America of early man's associations original 100 percent born in a given
of the Pleistocene. These include many with these mammals, as noted above, spring. Data for a typical large
grazing mammals which did not is most informative. Only the mam herbivore survivorship curve under
significantly compete with early man moth and, especially, the bison occur natural pr?dation come from a recent
for food resources. in great numbers as the victims of study of the wolves and moose of Isle
hunting activities of Paleo-Indians. Royale in Lake Superior (Mech 1966).
The number of genera (33) named by The absence, in most cases, of the Here, well over half (62%) of each
Martin (1967) as recently becoming remains of horse, camel, mastodon, or year's crop of young moose is taken by
extinct in North America by "over pronghorn from kill sites and their the wolves in that year before winter

166 American Scientist, Volume 58

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sets in. After that, the young adults
can better defend themselves and
rarely fall victim for the next several
years. Finally with increasing age
the moose are again unable to escape
the wolves.
In his work on the wolves of Mt.
McKinley, M?rie (1944) found the
bighorn sheep had a similar survivor
ship distribution, as based on ages of
collected skulls. Murie's figures did not
show as large a proportion of first-year
kills; but it is clear, as he notes, that
skulls of very young sheep would be
more apt to disintegrate or be eaten
than older skulls. With some cor
rection for this factor, the age distri
bution of these sheep agrees with that
for Mech's moose.
Other data which are less easily
graphed all indicate a massive car Fig. 1. Typical survivorship curve of large
nivore kill of each year's young, fol herbivore under natural pr?dation.
lowed by a plateau of little pr?dation,
and finally the gradual elimination of
the aging animals. In terms of the
yearly cycle, it is during the winter
that hunting man, if present, is
pressing hardest for largely the same
victims in lieu of significant vegetable
foods. Competition with man in this
season alone could be sufficient cause
for local disappearance of those car
nivores whose prey happened to be the
same as man's.
In the absence of the natural car
nivores, the herbivorous species would
have a survivorship curve which
follows mainly from human pr?dation.
This becomes a rather different pic
ture, especially after the development
of projectile weapons with which a
man can dispatch adult animals al
most as easily as young ones. Human
hunters may also, in many cases,
emphasize the young and aged game, Fig. 2. Typical survivorship curve of large
but generally not to the same degree herbivore under human pr?dation.
as most carnivores. Data on human
game kills tend to indicate a more
nearly equal take at all ages of the Carnivore kills. There is no leveling Other recent data on age distribu
prey species. If such pr?dation is off or plateau at the young adult tions of human kills include Friley on
severe enough, the surviving numbers stage. otter (1949a) and on beaver (1949b).
of reproducing adults may be no more These records are graded not by age
This chamois curve representing the in years but rather by degrees of
than when natural carnivores are
take of one hunter also shows the age
involved, but in many instances this is maturity; still they indicate the same
distribution of the population from curve of survivorship, with no plateau
not what happens. which they were taken according to at any stage. More data, especially on
Age distribition of prey populations availability. The age distribution was kills by primitive hunters, would be
under human pr?dation is clearly caused, in turn, by this and previous useful to illustrate this point.
shown in one case. Bourli?re (1956: p. hunting which also took animals at all
293) records the extensive take of ages with no emphasis on the young
chamois by one hunter in the Alps and and aged and no avoidance of young Some archeological data are also
Pyrenees. Here there is no killing of adults. The curve might be com available on the proportions of young
the youngest animals, but the rest of pleted by adding an estimate, probably individuals recovered from occupation
the survivorship curve is distinctly high, of 20 percent more as having and butchering sites of ancient man.
different from that resulting from died in their first year. Butzer (1964: p. 382) records 19 per

1970 March-Apr il 167

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 cent as being "juveniles" among the
reindeer taken at the Lake Mousterian
site of Salzgitter-Lebenstedt. Soergel
(1922, as cited by Butzer 1964: p. 388)
mentions proportions of immature in

3te |^^^'^^'~ ' ^ 'r':: ^ ' ^4^/^''^'

dividuals of numerous species of game
ranging from 25 to 35 percent in
various European Paleolithic sites. In
America, Hester (1967: p. 181) notes
that Paleo-Indians took game of all
ages with no particular emphasis.
While some of these figures indicate
a rather high mortality of young game
animals, they are all still far below the
over 50 percent which is indicated for
natural carnivores.

A comparison can now be made of the

two survivorship curves with natural Barth ,v * - : *y; 6 ~' .* ^ ; ? ^ >' m?. Ift^l?., - Yelt*
carnivore pr?dation (based on Mech
and Murie) and with human pr?dation
(based on Bourli?re, Friley, and Fig. 3. Superimposed survivorship curves showing differences between natural and human
others). The wide variation possible pr?dation.
in each of these curves could, in some
instances, negate the observations that 4. Other herbivores are exterminated did not apply to the mammoth will
follow. In general, however, the differ by pressure from the "favored prey" be discussed below.)
ence between human and other car through competition for such things as 4. Bison exterminated many other
nivore pr?dation practices tends to food under stress conditions. species which were in close ecological
cause a marked contrast in the age 5. Still other carnivores may next be competition. Thus the demise of such
distributions of the prey population. come extinct as their prey disappears, other plains herbivores as horse, ex
if they cannot depend on the expand tinct pronghorn, various cattle, camel,
Most conspicuous is the difference in ing type.
and perhaps also the mammoth.
numbers of surviving young adults.
Under human pr?dation these indi Of course, some rather close predator 5. The jaguar, presumably a predator
viduals, the major breeding stock, are prey relationships must already have on horse, camel, or cattle (or all of
far more numerous because their num been in existence for this sequence of them), then disappeared from the area.
bers were not so drastically depleted events to occur. If the predator which The above is probably a minimal de
when they were still immature. The man competes with has another major scription of events. Other species may
numbers of surviving older adults food supply during seasons of stress, it have been involved through ecological
might be greater under natural pr?da will not become extinct. If other com relationships which are not obvious
tion, but this would not generally be peting carnivores are taking significant to this writer. In environments other
enough to outweigh the breeding po numbers of the young of this game than the plains, Paleo-Indians might
tential of the younger ones. It can thus species, its population will not ex have had other game specializations
be seen that when natural predators pand. Even with only moderate with similar effects on still other species.
are replaced by hunting man the most predator-prey specializations the chain The extinction of the woodland masto
likely expectation is an increase in the of events outlined should occur to don suggests competition from other
population of the prey. If all else re some degree. animals which have survived, but
mains equal, this prey species ought there are no data on this comparable
then to exert considerable population In the case of the prominent faunal with that on plains Paleo-Indian
pressure against any competing species extinctions in North America, a activities.
in the area with similar ecological re reconstruction of the particular events,
quirements. according to this scheme, might have Just why the American mammoth dis
been the following: appeared remains a problem. Eco
A general picture of a series of bio 1. The first hunting by man with logical pressure from bison seems the
logical events can now be suggested : projectile weapons in North America most reasonable explanation, while
1. Human hunters enter an area and began about 12,000 years ago (Haynes climate change may have been a major
specialize in taking a particular species 1967), with emphasis on killing mam factor as well, and human pr?dation
of game animal. no more than a minor factor.
moths and especially bison.
2. The local carnivore (or carnivores) 2. The saber-toothed cats which de Some light may be thrown on the
specializing in the same prey are pended upon the same two species, reason for the American mammoth's
starved out during winter competition especially on the aged animals in extinction by a set of figures given by
and become extinct. winter, were competed out of exis Soergel in 1922 (quoted by Zeuner
tence.
3. With man taking proportionally 1963: p. 17) on the age distribution of
fewer of the young, the prey species 3. Bison greatly increased in numbers fossil elephants of Pleistocene Europe.
expands in numbers and presumably in because their young were no longer From Soergel's data two survivorship
territory. being killed by sabertooths. (That this curves can be drawn?one from first

168 American Scientist, Volume 58

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 ply. The aborigine's concentration
on kangaroos and wallabies might
have relieved other pr?dation pressure
and permitted these medium-sized
marsupials to exterminate other larger
species. More evidence is needed to
pursue the Australian situation any
further.

Agricultural Man
The third method by which man has
been able to exterminate species
without direct killing is by agricul
turally related practices. Three of
Martin's associations between the ad
vent of man and the disappearance of
big game animals are the more recent
cases of the West Indies, New Zealand,
and Madagascar (see also Hooijer
1967; Battistini and V?rin 1967; and
Fig. 4. Survivorship curves of Pleistocene European elephants under human pr?dation Walker 1967). In each case, the
(late), and presumed natural pr?dation (early).
human population at the time of ex
tinctions was of the Neolithic type,
with the subsistence economy based
interglacial fossils and the other from last stage, the Magdalenian, there was on agriculture. There is little doubt
fossils of third interglacial times. an almost total concentration on
that these people were responsible for
Zeuner assumes man was responsible reindeer as the game animal. One
the disappearance of numerous species,
for most of the kills in each group, but might conclude that man gradually
but not necessarily in the sense of
I suggest that sabertooths were the shifted his hunting emphasis solely
hunters destroying their prey.
earlier predator. It is an educated according to availability of game
guess that only a small difference species. It seems more likely that the There are several ways in which man,
would exist between the ability of increasing abundance of reindeer was since the Neolithic revolution, has up
man and of sabertooths to kill ele largely a result rather than a cause of set natural environments. Farming
phants; thus one might expect a man's concentration on this species. may remove local vegetation which
close similarity in the two survivor was a food supply for some animals.
ship curves. Yet there is a noticeable In discussing the Meiendorf site of Fencing will restrict some animals'
13,500 b.p., Butzer (1964 : p. 410) stated
difference, the earlier pr?dation pat movements as well as deny them access
this proposition indirectly: "There is
tern being more like the natural one, to the planted crops. Forest clearing
with a greater "take" of young no ready explanation why bison and will remove habitats as well as certain
animals and a slight tendency to level
woolly mammoth should be com food sources, substituting others.
pletely absent in the Hamburg area
off in most of the adult years. Draining swamps and irrigating dry
other than by reason of deliberate lands likewise alters vegetation and
Unlike most herbivores, elephants specialization on reindeer by pre environment. Stock raising introduces
begin breeding at a relatively late historic man." Perhaps this was not major competitors for certain food
age?about sixteen years. While there intended to mean that man's special resources. The physical presence of
is some separation between the two ization on one species was the cause of man with his activities and construc
curves, representing more elephants the absence of others in the area
tions can interfere with mating, nesting,
alive in early adulthood under hu (not just the site), but that is what
or maternal behavior, or simply in
man pr?dation, this is more than I am suggesting. duce animals to leave an area to avoid
canceled by the late onset of reproduc
tion. These rather scanty data suggest
In some European regions a corre such disturbances.
that elephants, unlike the bison, sponding emphasis on horse hunting The possibility that some of these
seems to have occurred. If we could
would not increase in population after agricultural activities have caused
human pr?dation replaced that of identify the natural predators involved exterminations has been pointed out
saber-tooth cats. we might find a similar situation. The by Guilday (1967: pp. 122, 126, 138)
same events may have occurred in
for much of the megafauna in North
In Europe a chain of events similar to Australia, but adequate archeological
Africa and the Middle East, and by
that conjectured for North America data are lacking. Gill (1955), in Hester (1967: p. 178) for North
seems to have occurred, though with reviewing the situation, was able to
America since European coloniza
less drastic consequences in terms of conclude only that man could not tion.
numbers of kinds of animals exter have been the direct cause of the
minated. Throughout the Upper Paleo extinctions. Here, annual variation In addition to his great ability to affect
lithic there was an increase in emphasis in rainfall, rather than winter cold, is native fauna indirectly, the agricul
on reindeer hunting in Western Eu the probable source of the stress turalist differs from the hunter in his
rope and a decrease in hunting mam season during which one species elim lack of dependence upon that fauna.
moth and woolly rhinoceros. By the inates another by taking its food sup The extinction of numerous large

1970 March-April 169

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native herbivores will have little or no broad category will have an exchange Haynes, C. Vance, Jr. 1967. Carbon-14
dates and early man in the New World, in
ill effect on the farmer who depends on value based, in part, upon their
Martin and Wright, pp. 267-86.
his crops and/or herds; he may even scarcity. Hunters may thus concen
trate on certain animals not for their Heizer, Robert F. 1955. Primitive man as
welcome their disappearance. The an ?cologie factor, in Kroeber Anthropological
hunter, on the other hand, would own use but for sale or trade with Society Papers, 13: 1-31.
perish without them. agriculturally based peoples. In these Hester, James A. 1967. The agency of man
cases a decline in the supply of a in animal extinctions, in Martin and
While many writers have missed the particular animal may raise its price Wright, pp. 169-92.
point by emphasizing the Paleo and serve to increase pr?dation pres Hooijer, D. A. 1967. Pleistocene vertebrates
Indian's dependence on big game sure. Just as the African elephant of the Netherlands Antilles, in Martin and
hunting to account for his supposed now has the added threat of ivory Wright, pp. 399-406.
destructiveness, Edwards (1967: pp. hunters, it may be possible that the Jelinek, Arthur J. 1967. Man's role in the
147-48) realized that only by re last Siberian mammoth was killed for extinction of Pleistocene faunas, in Martin
acquiring plant-gathering habits the Chinese ivory market. and Wright, pp. 193-200.
would primitive man have been able Kowalski, Kazimierz. 1967. The Pleistocene
to afford the luxury of exterminating Let me add that the above arguments extinction of mammals in Europe, in
Martin and Wright, pp. 349-64.
his game. This, of course, does not are not proposed as the entire explana
Kurten, Bj?rn. 1968. Pleistocene Mammals of
explain how such extermination was tion of megafaunal extinctions. Cli
Europe. Chicago: Aldine.
possible on a hunting and gathering mate change cannot be ignored, be Leopold, Estella B. 1967. Late-Cenozoic
cause the combination of an unusually
level, but it does illustrate the point patterns of plant extinction, in Martin and
that man can destroy only what he abrupt end to the last glaciation with Wright, pp. 203-46.
does not depend upon, without de the subsequent altithermal warming Martin, Paul S. 1958. Pleistocene ecology
stroying himself in the process. may have been unique in the Pleisto and biogeography of North America, in
cene. Human agency, by the means Zoogeography, G. L. Hubbs, ed., American
The recent destruction of megafauna suggested here, is proposed to account Association for the Advancement of Sci
in New Zealand, Madagascar, and the for extinctions only where climate ence, Publication 51, pp. 375-420.
West Indies followed from the activi - 1967. Prehistoric overkill, in Martin
change is an inadequate explanation.
ties of agricultural man rather than of and Wright, pp. 75-120.
hunters. This points up a meaning Martin, Paul S., and John E. Guilday.
ful dichotomy of human societies in 1967. A Bestiary for Pleistocene biologists,
terms of their effect on the environ in Martin and Wright, pp. 1-62.
Martin, Paul S., and Herbert E. Wright,
ment. The contrast between "pre Bibliography Jr., eds. 1967. Pleistocene Extinctions, vol. 6,
historic" and "modern" man in this Battistini, R., and P. V?rin. 1967. Eco
Proceedings of the Eighth Congress of the
connection is not of much value. logical changes in Protohistoric Mada International Association for Quaternary
The "modern" Plains Indians and gascar, in Martin and Wright, pp. 407-24. Research. New Haven: Yale University
Press.
African Bushmen have had a very Bourliere, Fran?ois. 1956. The Natural
History of Mammals. New York : Knopf. Mason, R. J. 1962. The Paleo-Indian tradi
different effect on their environments
Butzer, Karl W. 1964. Environment and tion in Eastern North America, in Current
than did the simple grain farmers of Archeology. Chicago: Aldine. Anthropology, 3: 227-78.
the Eastern Mediterranean some eight Colbert, Edwin H. 1942. The association of Mech, L. David. 1966. The Wolves of Isle
or ten thousand years ago. Agricul man with extinct mammals in the Western Royale, Fauna of the National Parks of the
turally based societies differ from one hemisphere, in Proceedings, Eighth American United States, Fauna Series 7. U.S.
another only in the degree to which Scientific Congress, 2: 27. Government Printing Office.
they disturb the natural environment, Cornwall, I. W. 1968. Prehistoric Animals Murie, A. 1944. The Wolves of Mount
and Their Hunters. London: Faber & Faber.
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Edwards, William Ellis. 1967. The Late the U.S., Fauna Series 5. U.S. Govt.
ing and gathering societies. Pleistocene extinction and diminution in Printing Office.
size of many mammalian species, in Newell, Norman D. 1963. Crises in the
There are some additional peculiarities Martin and Wright, pp. 141-54. history of life, in Human Variations and
of agricultural societies, in terms of Eiseley, Loren C. 1943a. Archaeological Origins, W. S. Laughlin and R. H. Os
size of population and commerce, observations on the problem of post borne, eds. San Francisco: Freeman.
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that they could have exterminated 8: 209-17. Romer, Alfred S. 1933. Pleistocene verte
- 1943b. Did the Folsom bison survive brates and their bearing on the problem of
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in Canada? in Scientific Monthly, 56: 468-72. human antiquity in North America, in
tion. Given a farming subsistence
Flint, Richard F. 1957. Glacial and Pleisto The American Aborigines, ed. by D. Jenness,
base, almost any area will support and pp. 76-77.
cene Geology. New York: Wiley.
feed a population ten to a hundred Simpson, George G. 1965. The Geography of
Friley, C. E., Jr. 1949a. Age determination,
times larger than it could maintain by use of the baculum in the river otter, Evolution. Philadelphia: Chilton Books.
by hunting and gathering alone. Slaughter, Bob H. 1967. Animal ranges as a
Lutra c. canadensis, in Journal of Mammalogy,
With such numbers available, espe 30: 102-10. clue to Late-Pleistocene extinction, in
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170 American Scientist, Volume 58

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