Professional Documents
Culture Documents
American Association for the Advancement of Science is collaborating with JSTOR to digitize, preserve and
extend access to Science.
http://www.jstor.org
Megafaunal Biomass
Paul S. Martin
author
is
professor
of
geosciences,
Uni-
Table 1. Large mammal biomass in some African parks and game reserves [from Bourliere
and Hadley (32)]. [Courtesy of Annual Reviews, Inc., Palo Alto, Calif.]
Biomass
(metric
tons per
square
kilometer)
Biomass
(animal
units per
square
mile)
Location
Habitat
Number
of
species
TarangireGame Reserve,
Tanzania
Kafue National Park,
Zambia
Kagera National Park,
eastern Rwanda
Nairobi National Park,
Kenya
Serengeti National Park,
Tanzania
Queen Elizabeth National
Park, western Uganda
Queen Elizabeth National
Park, western Uganda
Albert National Park,
northern Kivu
14
1.1
Tree savanna
19
1.3
Acacia savanna
12
3.3
18
Open savanna
17
5.7
32
15
6.3
36
11
12
68
11
27.8-31.5
158-179
11
23.6-24.8
134-141
The alternate view, that the American large mammal biomass was in
eclipse during the late glacial (12),
cannot be tested quantitatively on the
basis of fossils alone. Bones do not provide reliable estimates of past biomass
(13). But the great numbers of mastodon, mammoth, extinct horse, camel,
and bovid bones found in late-glacial
sediments hardly suggest scarcity. Evidence that the Late Pleistocene megafauna was declining in numbers and
diversity before 12,000 years ago, as
Kurten found in Late Paleolithic sites
of the Old World (14), is lacking in the
New World (11).
Table 2. Simulated values for New World population growth.* In example 1, 104 people
reached Edmonton 11,500 years ago. They double in numbers every 20 years until limited
by their southward migration rate, 16 kilometers per year. Population growth fills a sector
of 90? and is concentrated along the arc ("front") through a depth of 160 kilometers.
The density on the front is 0.4 person per square kilometer and behind the front is 0.04
person per square kilometer. Example 2 is the same as example 1 except that the population
doubles every 30 years. Example 3 is the same as example 1 except that the migration rate is 8
kilometers per year. Example 4 is the same as example 1 except that the migration rate is 25
kilometers per year. Example 5 is the same as example 1 except that the front is 80 kilometers
deep. Example 6 is the same as example I except that the front is 240 kilometers deep.
Example 7 is the same as example 1 except that the front is 0.02 person per square kilometer.
Example
1
2
3
4
5
6
7
8,000
8,000
8,000
8,000
2,800
18,000
8,000
Distance
(km)
184
299
159
199
172
193
180
393
583
207
583
457
349
368
Population
61,000
102,000
26,000
102,000
40,000
81,000
53,000
Population
590,000
590,000
590,000
590,000
435,000
750,000
450,000
Local faunal
= extinction
Modeling Overkill
The impact of the hunters is best 1
visualized if one considersa representa0.4
tive area on their front. If a sizable
biomass,say, 50 animal units per square
Human
mile, were exposed for 10 years to
population
0.2
hunterswhose density is one person per
density per
square mile on the average, what resquare
moval rates would be necessary to rekilometer
duce the fauna? The fraction of the
standing crop of moose available an4
1
/
lU
nually to wolves on Isle Royale is 18
older
and
percent (20); mainly
Time (years)
young
animals are taken. For animals larger
of the front showing theoretical changes in human population density.
Fig. 1.
than moose, an annual removal rate of At any Passage
one point, the big game hunters and the extinct animals coexisted for no
20 percent of the biomass attributable more than 10 years. Poor paleontological visibility of kill sites is thus inevitable.
9 MARCH 1973
971
reviewed the historicrecordsof extraordinary meat consumption and occasional extreme waste among the Plains
Indians.
Unless one insists on believing that
Paleolithic invaders lost enthusiasmfor
the hunt and rapidly became vegetarians by choice as they moved south
from Beringia, or that they knew and
practiced a sophisticated, sustained
yield harvest of their prey, one would
have no difficultyin predictingthe swift
exterminationof the more conspicuous
native American large mammals. I do
not discountthe possibilityof disruptive
side effects, perhaps caused by the introduction of dogs and the destruction
of habitat by man-made fires. But a
very large biomass, even the 2.3 X 108
metric tons of domestic animals now
ranging the continent, could be overkilled within 1000 years by a human
population never exceeding 106. We
need only assume that a relatively innocent prey was suddenly exposed to
a new and thoroughly superior predator, a hunter who preferred killing
and persistedin killing animals as long
as they were available (21).
With the extinction of all but the
10,800 N
years ago
10,700 years
ago
ago
10,500 years
ago
'The Front'
40 people per 100 km2
Lii
:3
Scale
0
kilometers
Scale
0
iI
500
I
1000
1600
kilometers
Fig. 2 (left). Sweep of the front through North America. As local extinction occurs, the hunter moves on. 3 Fig. 3 (right).
Sweep of the front through South America. Local extinction accompanies passage of the front. (Figures 2 and are not drawn
to scale.)
972
~~~SCIENCE,
~~~~~~~~~~~~~~~~~~972
VOL. 179
Summary
I propose a new scenario for the
discovery of America. By analogy with
other successful animal invasions, one
may assume that the discovery of the
New World triggered a human population explosion. The invading hunters
attained their highest population density along a front that swept from
Canada to the Gulf of Mexico in 350
years, and on to the tip of South America in roughly 1000 years. A sharp
drop in human population soon fol-
tion is not foolproof and because bone contamination is not always avoidable, we may
expect a steady increase in postglacial dates
of the sort which misled me years ago [P. S.
Martin, in Zoogeography, C. Hubbs, Ed.
(Publication No. 51, AAAS, Washington,
D.C., 1958), p. 397]. Admittedly, there is no
theoretical reason why a herd of mastodons,
horses, or groLundsloths could not have survived in some small refuge until 8COO or
even 4000 years ago. But in the past two
decades, concordant stratigraphic, palynological, archeological, and radiocarbon evidence to demonstrate beyond doubt the postglacial survival of an extinct large mammal
has been confined to extinct species of Bison
[see S. T. Shay, Publ. Minn. Hist. Soc. (1971);
J. B. Wheat, Sci. Amer. 216, 44 (January
1967); Amer. Antiquity 37 (part 2) (No. 1)
(1972); D. S. Dibble and D. Lorrain, Tex.
Ment. Mus. Misc. Pap. No. 1 (1968)]. No
evidence of similar quality has been mustered
to show that mammoths, mastodons, or any
of the other 29 genera of extinct large mammals of North America were alive 10,000
years ago. The coincidence in time between
massive extinction and the first arrival of
big game hunters cannot be ignored.
4. M. I. Budyko, Sov. Geogr. Rev. Transl. 8
(No. 10), 783 (1967).
5. According to R. F. Flint [Glacial and Quaternary Geology (Wiley, New York, 1971), p.
778], "The argument most frequently advanced against the hypothesis of human agency
is that in no territory was man sufficiently
numerous to destroy the large numbers of
animals that became extinct."
6. Apart from postglacial records of extinct
species of Bison, very few kill sites have
been discovered. J. J. Hester [in Pleistocene
Extinctions, the Search for a Cause, P. S.
Martin and H. E. Wright, Jr., Eds. (Yale
Univ. Press, New Haven, Conn., 1967), p.
169], A. J. Jellinek (ibid., p. 193), and G. S.
Krantz [Amer. Sci. 58, 164 (1970)] have all
raised this point as a counterargument to
overkill.
7. The North American megafauna that I believe
disappeared at the time of the hunters includes
the following genera: Nothrotherium, MegaEremotheritun, and Paramylodon
lonyx,
(ground sloths); Brachyostracon and BoreoCastoroides
stracon (glyptodonts);
(giant
beaver); Hydrochoerus and Neochoerus (extinct capybaras); Arctodus and Treinarctos
(bears); Smilodon and Dinobastis (saber-tooth
cats); Mamnutt (mastodon); Maimmuthus
(mammoth); Equtts (horse); Tapirus (tapir);
Platygonits and Mylohyus (peccaries); Camtelops and Tattipolama (camelids); Cervalces and
Sangaimona (cervids); Caprotneryx and Tetrameryx (extinct pronghorns); Bos and Saiga
(Asian antelope); and Bootherittm, Symbos,
Euceratherittm, and Preptoceras (bovids).
8. C. V. Haynes, in Pleistocene and Recent EnIironments of the Central Great Plains, W.
Dort, Jr., and J. K. Jones, Jr., Eds. (University of Kansas Department of Geology Special
Publication No. 3, Lawrence, 1971), p. 77.
9. A. Dreimanis [Ohio J. Sci. 68, 257 (1968)]
estimates that there are more than 600 mastodon occurrences in northeastern North
America. If we suppose that 500 of these were
of late-glacial age and assign an equal probability of death, burial, and discovery to each
in the time span from 10,500 to 15,500 years
ago, then an average of ten may be expected
for any given century and one for any decade.
I assume that in any one region local extinction was swift. The elephants and their hunters
were associated for no more than a decade.
Even if the temporal overlap between the
elephants and their hunters were as much
as 100 years, and if half (five) of the finds
represent animals killed by hunters, it is
clear that the probability of the field evidence
actually being detected and appreciated by
the discoverers of the bones is small. Had
the hypothetical mastodon kill sites been
located on the uplands rather than in bogs
or on lake shores, the probability of discovery becomes smaller still. My pessimistic
appraisal should not deter those engaged in
the search for more kill sites. It should refute
the view that extinction by overkill would
yield abundant fossil evidence.
10. One animal unit can be used as a standard
for paleoecological comparison in the sense
range managers have used it for comparing
974
22.
23.
24.
25.
26.
the
Amrerican
Indian
(Pan-American
Rosenthal,
Experinmenter
Effects
in
Be-