The Empty Forest

Many large animals are already ecologically extinct in vast areas

of neotropical forest where the vegetation still appears intact

Kent H. Redford

he world conservationcom- With few exceptions, researchers

munity has focused much of We must not let a have concentratedon directalteration
its attention on the plight of of vegetation,not discussingthe ways
tropical forests. Many authors have forest full of trees fool in which human activities have af-
lamentedthe loss of forest cover and fected the animals of tropical forest
the destructionof the forestand spec- us into believing all ecosystems. In this article, I expand
ulated on the extent of the tropical is well the focus to include defaunationof
forest left intact. Throughoutthe dis- tropical forests, concentratingon the
cussion, tall, majestic, tropical trees forests of the Amazon basin, and I
are used as a symbol for the complete show that the long-termpreservation
set of animaland plant speciesfound stroying the forest from the outside. of tropical forest vegetation will not
in tropicalforests.Treesare also being In any case, ecologists looked for be possible if the forest fauna is not
used by some conservationbiologists, study sites that would allow for ex- also preserved.
parkplanners,and othersto represent aminationof "natural"processesun-
the entiretropicalforestbiota and as a contaminated by anthropogenic ef- Indirectdefaunation
measureof conservationworth. fects. Data from botany,archaeology,
The presenceof soaring,buttressed and anthropologycollected in many Humans can devastate a fauna by
tropical trees, however, does not parts of the world are showing, how- indirector direct means. Indirectde-
guarantee the presence of resident ever, that anthropogeniceffects are faunationis the destructionof a fauna
fauna. Often trees remainin a forest ubiquitousand that the sought-after through human activity not aimed
that humanactivitieshave emptiedof virgin habitat may not exist. Flenley specificallyat animals.In tropicalfor-
many of its large animals. The ab- (1979), for example,has documented ests, habitat destructionis the most
sence of these animals has profound widespreadhumaneffectson tropical common of these practices-not sur-
implications,one of which is that a forests throughoutthe equatorialre- prisingly,many forest animalscannot
forest can be destroyed by humans gions. survive without forest. A less-often-
from within as well as from without. The relativelyrecent arrivalof hu- consideredtype of habitatdestruction
Until recently,human influenceon mans in the western hemispherehas occurswhen animalsare absentfrom
tropical forests through such activi- not lessened the overall impact our an area of otherwiseexcellenthabitat
ties as burning,swidden agriculture, species has had on neotropical for- becausesome criticalarea elsewhere,
and hunting was regardedby ecolo- ests. From the forests of Mexico such as a nesting beach, was de-
gists as of such low impactthat it was through Panama, and the montane stroyed.This problemaffectsanimals
negligible,as importantbut confined forests of Colombia to Ecuador,sci- including migratory birds, beach-
to areas of human settlement,or as entists have documentedthe ways in nesting turtles,and white-lippedpec-
confined to rapacious colonizers de- whichpre-Columbianhumansaltered cary (Tayassupecari)herds.
the presence,extent, and structureof There are many other types of in-
forests. The forests of the Amazon direct defaunation.One of the most
Kent H. Redford is the director of the basinwere also extensivelyalteredby important is probably the effect of
Programfor Studiesin Tropical Conser- humanactivities.In fact, Balee(1989) forest-extraction activities by hu-
vation and an associate professorin the
Center for Latin American Studies and has recently suggested that at least mans. For example, logging can re-
Departmentof Wildlife and Range Sci-
11.8% of the terrafirmeforestsof the move fruit-bearingtrees and destroy
ence, Universityof Florida, Gainesville, Brazilian Amazon, almost 400,000 nesting and other criticalareas.
FL 32611. ? 1992 AmericanInstituteof km2, show continuingeffects of past Less obvious are the effects that
BiologicalSciences. human interference. stem from the much-publicizedex-

412 BioScienceVol. 42 No. 6

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 I
tractionof forest fruits and nuts. Al- INDIANS COLONISTS
most without exception, the fruits
collectedfor sale are fruits also eaten
by largebirdsand mammals(Redford Cebus apella
et al. in press).However,the extent to
which animalspeciescan survivewith
reducedavailabilityof fruitsand nuts 4B
sciurids
is unknown.
A recent article by Vasquez and
Gentry (1989) documentedthe huge Tayassu pecari
numberof fruits from wild trees col-
lected by humans for sale in Amazo-
Agouti paca
nian markets.One of the majorfruits
in this marketwas that from Mauritia
palms-the only food of the macaw Dasypus novemcinctus i
Ara manilata (Roth 1984) and the I I
most importantfruitin the diet of the
tapir (Bodmer1990). The enormous Tayassu tajacu
quantities of Brazil nuts removed
from a patch of tropical forest un- Dasyprocta &
doubtedlyaffectthe animalsthat oth- Myoprocta (d)
erwise would have fed on those nuts.
The extractive activities also remove Alouatta spp.
nutrientsfrom the ecosystem.
Indirectdefaunationcan also take
Ateles spp.
U
place through the effects of subsis-
tenceor commercialhuntingand fish-
ing that remove potential prey from Cebus spp.
tropical forests, thereby affecting
predators, scavengers, and the ani-
mals that depend on them (Thiollay Tamandua spp. I
1984). As Emmons (1987) and Jor- eE
genson and Redford (in press) have I
pointed out, every majorprey species
of the jaguaris intensivelyhunted by
Mazama spp.
h I
humans.In these cases, it seems clear Bradypus tridactyla
I
that the removalfrom tropicalforests
of food for human consumption re-
;CC
duces the capacityof those forests to Tapirus terrestris
supportmany animals. Xm-
Finally,manyof the by-productsof
modern human activities are impor- Figure1. Importanceof mammalsto contemporaryIndianand colonist hunters(only
tant contributorsto indirectdefauna- consideredwere those species found in a minimumof five Indian studies and three
coloniststudies).Barsdenotethe numberof individualsof that taxon killedper hunter
tion. These byproductsinclude mer-
per year.To give an idea of scale,therewere approximately2.5 individualCebusapella
cury and sediment contaminationof monkeys killed by Indiansper consumerper year and approximately0.05 Tapirus.
fish (Martinelliet al. 1988); smoke, (Data from Redfordand Robinson 1987.)
which especially affects pollinators
I
(Lovejoyet al. 1984); and an increase
of edge habitatin the forest (Malcolm vided into two categories:subsistence have lived for decadesin tropicalfor-
1991). huntingand commercialhunting. ests but have recourse to domestic
animals.It is also of lesserimportance
Direct defaunation Subsistencehunting.In manypartsof to colonists recentlyarrivedto Ama-
the world, wildlife serves as a major zonian forests, who frequently are
The indirecteffectsof human activity source of food for local peoples. In unfamiliarwith huntingand have ac-
on defaunation, important as they LatinAmerica,game is a vital protein cess to other sourcesof meat.
are, have arisen chiefly in recent dec- and fat source to many groups living A wide varietyof wildlifeis hunted
ades. The effects of direct defauna- outside of urban areas. As a general for food by humans. The Maraca
tion, the deliberate killing of animals, rule, wildlife is most important to Indians of Colombia, for example,
has a much longer history in the Am- Indian groups that depend on game take at least 51 species of birds, in-
azon forests-a history that coincides meat for subsistence. It is of lesser cluding 10 species of hummingbirds
with the presence of humans in the importance,thoughstill important,to (Ruddle 1970). Hunters generally
area. Direct defaunation can be di- settlers of European descent who take more mammals than birds and

June 1992 413

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 INDIANS COLONISTS and has shifted from group to group
as market demand and availability
have changed (Figure3; c.f. Redford
and Robinson 1991). Even before
Penelope spp. I Europeans arrived in the Americas,
animals and their products were
Ramphastos spp. traded. Among the Incas, adult
caiman and anacondas were trans-
ported from the Amazonian low-
Crax spp. lands up to the Andeancity of Cuzco
th for use in menageries (Lathrap
1975). Nonetheless, trade in wildlife
Psophia spp. I did not assume major proportions
4b until Europeansarrived.
EDIBLE PRODUCTS. As early as the
Crypturellus spp.
k seventeenthcentury, the commercial
harvestingof manatees for meat be-
Amazona spp. I gan (Redford and Robinson 1991).
Until the mid-1900s, the averagean-
nual catch of these Trichechusinun-
Ara spp. I guis in the Amazon was at least sev-
eral thousand animals. Two other
Mitu spp. I importantcommercializedsourcesof
meat are caiman and river turtles.
Caimanof severalspecies,but princi-
Figure 2. Importanceof birds to contemporaryIndian and colonist hunters (only
consideredwere those species found in a minimumof five Indian studies and two pally of the genus Caiman,have been,
coloniststudies).Barsdenotethe numberof individualsof that taxon killedper hunter and still are, an importantsource of
per year. To give an idea of scale, there were approximately0.9 individualPenelope meat in some areas of the Amazon
guanskilledby Indiansper consumerperyearand approximately0.09 Ara. (Datafrom basin, with the currenttrade in meat
Redfordand Robinson 1987.) estimated at 21,500 to 32,000 ani-
mals annually.
The first Europeanto navigatethe
morebirdsthan reptiles(Redfordand I have estimated the number of Amazon River found many Indian
Robinson 1987). mammals killed in one year by the villageswith hundredsof pennedtur-
Throughout Amazonia and Latin rural population of AmazonianBra- tles (mostly Podocnemis expansa).
America, indigenous hunters usually zil. In 1980, there were an estimated Despite ferocious exploitation for
kill only a few of the many types of 2,847,000 people living outside of meat and eggs, female P. expansa
mammals and birds present. Of the cities in an area of 3,581,180 km2 continued to gather near nesting
mammals, monkeys, peccaries, deer, (FIBGE1982). I multipliedthis num- beaches and be plentiful enough
armadillos, and large rodents like berof consumersby the per capitaper through the 1850s to impede river
paca and capybara are frequently annum consumption values derived trafficon the Madeira.
hunted; and, of the birds, the most fromstudiesof colonisthunting(Red- Turtle eggs have been heavily ex-
common prey are guans and curas- ford and Robinson1987). The result- ploited for industrialand nutritional
sows, toucans, trumpeters, and ma- ing figure,14 millionindividualmam- purposes. In the Amazon basin, the
caws. The number of species hunted mals killed each year, suggests the eggs of P. expansa were so abundant
by nonnative peoples is even more staggeringextent of subsistencehunt- and in such great demand that an
restricted. Figures 1 and 2 show the ing. Adding birds and reptiles, the industry developed to process them.
differences in the range of mammal numberof game animals killed each Oil from the eggs was used for cook-
and bird species taken by Indians and year in Amazonian Brazil probably ing and lighting, and as early as the
colonists. reaches19 millionanimals.The num- eighteenthcentury royal decree con-
The numbers of animals taken by ber of animals fatally wounded or trolled the lucrativeharvestin Brazil.
subsistence hunters can be large. In killed could reach57 million animals In 1719, 192,000 pounds of oil,
less than a year, the 230 inhabitants a year. equaling approximately 24 million
of three Waorani villages in Ecuador eggs, were produced from the upper
killed 3165 mammals, birds, and rep- Commercial hunting. The second Amazon;as late as the 1860s, at least
tiles (Yost and Kelley 1983). This major cause of direct defaunationis 48 million eggs yearlywere harvested
total included 562 woolly monkeys commercial hunting. The killing of to supply the industry. In many ar-
(Lagothrix lagothricha), 313 Cuvier's animals in Amazon forests by Euro- eas, giant riverturtles have been vir-
toucan (Ramphastos cuvieri), and peans for commercial purposes has tually eliminated, and there is still
152 white-lipped peccary. Not all been going on since soon after their heavy predation on their eggs wher-
subsistence hunting is of this inten- discoveryof the continent.This trade ever and whenever they can be
sity. has involved many different species found.

414 BioScience Vol. 42 No. 6

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 ,

Although no longer available on Caiman sclerops skins. In the 1960s, apparentlyin response
the scale once observed,game is still MAMMALIANSKINS.Just as was the to overexploitationof jaguarand the
readily obtained in many local mar- case with leather,large-scalecommer- concomitantdecreasein numbers,the
kets. Castro et al. (1975-1976) re- cialization of skins was brought to cat trade shifted to the smaller spe-
portedthe meat of 24 speciesof wild- Latin America by the Europeans. cies.
life, includingsix speciesof primates, Both tradesare directedat the luxury The period between the end of the
for sale in the markets of Iquitos, marketsin Europe,Japan,and North Second World War and the early
Peru. They estimate that the inhabit- America.The skin trade has always 1970s was the golden era of the trade
ants of the Peruvian departmentof focused on a relatively few species. in skins originatingfrom the Amazon
Loreto, which includes the city of The trade in skins originatingin the Basin (McGrath 1986). In the 20
Iquitos, kill 370,000 monkeys annu- Amazon basin has been well docu- years since 1946, the Amazon River
ally for consumptionand sale. mentedand has concentratedon giant port of Iquitos, Peru, exported
The differencebetweencommercial otter (Pteronurabrasiliensis),riverot- 22,644 giant otter skins, 90,574 river
and subsistencehunting is becoming ter (Lutra longicaudis),jaguar (Pan- otter skins, 12,704 jaguarskins, and
increasinglyblurred.For example, in thera onca), and ocelot (Felisparda- 138,102 ocelot skins. The value of
a study of the ungulateharvestin one lis). Much smaller numbers of Felis skins caused people to move into
watershednear Iquitos,Peru,lumber- wiedii and Felis tigrinaskins are also sparsely inhabited areas and devote
men hunting to supply their camps traded. tremendous effort to commercial
with food accounted for 51% of the The trade in cat skins began with hunting. For example, one family of
ungulate harvest, illegal commercial jaguarsat the end of the last century. hunters in the EcuadorianAmazon
hunters for 11%, and subsistence
huntersfor only 38% (Bodmeret al.
1988). Date
LEATHER. The earliest European 1500
;V 16Rn
I.IV 1700
I
,-~V 1R80
Ivv~ 1Ivvv 00 1950 1 990
v 1,.v Iv

commercial exploitation of wildlife

for nonedibleproductswas the use of Product

leather, particularly that made from MANATEES meat ~~~~~~~~~~~~~~~~... ~

ii~?;.;~.
**
????1?.1;.,...r?~~
,
,.... .....
.. .. ............ .... \,,,.,,.*--
Z
..*ss*.*s
.................s*
+v

deerhides. Most of the recentmarket hides ............

. . ........ . .........????? ?:?.?:?......................
???????? .;l
?????

for leatherhas been luxuryitems such meat ;;;.,

.;:2,- ;;-; "'-*
'v-
s~~~~~~~~~~~~~~~.:.:-.:..... . ;_s_g
as purses,gloves, and expensiveshoes TURTLES eggs ','.' .' . .....' -
:'.,
,r
.''=
.......
.-.' ;. ' ' -''
' ' -

oil
and overcoats.The principalanimals
killed for this trade are peccaries, skins
CAIMAN meat
capybara,and various species of rep- oil
tiles.
OTHER REPTILES
Peccary leather has always been Caiman lizard I...-:, ...::.>.,,....v.af.....
...-.
popular, particularlyin Europe and
-::
-.-: .: ./,:..,...:
.........
,.,..s
Iguana, Anaconda, skins
Japan.Between1946 and 1966, more Boa
?a~:::~~~~~~~~~~~iiQ
than 2 million collared peccary (Ta- ' ' . :, . : ..:>
:,,.......:

skins and meat

yassu tajacu) 800,000 CAPYBARA
leather
white-lipped peccary skins were ex-
ported from Iquitos, Peru, alone. DEER skins
Capybaraare anothersourceof high- meat
quality leather. Between 1960 and ..
ts:f ->.
..?..
-::::::::::: ..
....:
1969, almost 500,000 capybaraskins PECCARY skins
wereexportedfromthe BrazilianAm- meat
azon. Tradein these speciescontinues
today, although at substantiallyre- OTTERS skins
duced levels.
The most importantwildlife in the
skins
leather industry at present are the CATS
reptiles, principallythe crocodilians.
During the peak of the trade in the PRIMATES biomedical
1950s and 1960s, five to ten million trade
crocodilian skins were reported as ..,
,a''as''
. ,,,,,. .,
^ ,,
fS:::,y.........:S ...............
traded annuallyworldwide, with the BIRDS plumes
X>-?:?:?~?~?~?:?~;?~?~,:?:
* . * f8">2' ..- ' -. * : , ?i:-::.. ......... .......
...... .........'. . ...-...

actual figure probably much higher. X __~::~~: _ W i =

The extent of the trade is staggering:
for example, in Venezuela during PARROTS/MACAWS pets

1930 and 1931, 3000-4000 caiman

skins were being sold daily, and be-
tween 1951 and 1980 Colombia le- Figure3. Commercial of faunain theAmazonsincethetimeof European
exploitation
gally exported almost 12 million discovery.

June 1992 415

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 100 I ... . .*..... . . .**..*@@@@|@@...............
...... . . . . .

............
. . . . .. . .. . .. .

........
.. . .. . .. I...................................
.................................
................ ous ports of entry. Fromthe perspec-
............... *
?.................. .....
-....................... ..... .......... ........ .
................................... ........
........................... ............ ......
.... ........ ..... ... ...
tive of an ecologist, however,what is
.............

-.-,............... .... ...-.. ...-.....-..

of interest is the total number of
..- .. .....-.-.-
...................................
...................................
.......................,.....................
................................... ~ .:,~...
.o........,.....-. ....-..- .. ... .-.......,.:%.......!
... .....- ....' ....-,,., -...... individualanimalsremovedfromeco-
..., .......
90 -
.. . . . . . . .
...................................
.....-............. iiiiiiilll ......................................
....
...........
@*.. .. ..
...................................
.....................
,.-,.......,...
.. ..
.............
.. .. .. . . . . ..
. ..... ... . . .. ,-
......................... systems in the course of commercial
...................................
l .......... ............ . . . . . .
.........
.....
.....................
***...
'.-..,.-.,.........,.....,
..........
..
..... ..
.........
.....
.......
..
.. ... .............
..
...-...-...
..
. ...
..
. ...
..
...
..
-..
trade. For each animal that makes it
....,,
.. .
...
into the import/export trade, many
...................................
:::::::::...
. . ........... .. .. . ..........
.....:::::.:
..................
...................................
....................................
.,-.,.........
....................................
....................................
.....
@.. *....
.......
**................
.. others are killed.
... . ......
^

80 - , . ..........
........ . ..........
.....
.. .. .. ........... . . ..... .
?.-..-..-.-...--,....-.......-.................-.-.....
..
............... .................
. .
.. ..............
.......... ......
...........
The number of animals actually
.........
.. . ..........
..?....................-...-.-.-.
...........
.........................
.. .. ..-........
. . .
..
...........
...........
......-....
. . .......... i:::::iiiii Nogae i:i:i:i:*i:i:i:i:*
...........
. ....................................
....................................
.............
i:i*:i* Zkilled is unknown, but from the
*@@.
. . . . .

lim-
.

. . . . .. ..........
..... . ..... . . . . . . .,. . .. . ,.. . .. . . ..
................................... ... .
................................... ................-.-.-.-.-.-..-.....
...................................
...................................
................................... ........... .... Non-game.
ited data it seems reasonable to esti-
............-..-.-.--...................................
........
...................................
....................................
,...................................
...................................
..
. .. . . . . ...birds.....
::::::::::*:::*::-::*::-::*::-::-::|::@::@::::::::::|:-::|::@::-:::::::
...........
.
.... .. . . .
mate that for each animal enteredas
...........
::::::::::;:::::::No-a
::::::::::::::::: e::::::::::::::::::::
70 - ...................................
...................................
...................................
.........................................
...................................
....................................
....................................
~~~~~~~................-.-.....
an export statistic approximately
................................... ':':':':':':':':':':':':':':':':':':':':':':-:':':'::-:':-:-:':O:':':-:
.....
...................................
...................................
...................................
No-ampaide()
....
...................................
....................................
................................
.. ..... .... .... .....
three additional individuals die.
..... .....

...
...................................
...................................
....................................
.................................
....................................
...................................
...........
....................................
........... .......... These animals may be wounded by
,...................................
...................................
...................................
. .....................................
.. . ... -..-.-.-*.*..-.........
... ... ....
...........................................
... .......
....
....................................
...................................
... ...
.... .... .......... ....... the hunter or the hunter's dogs or
l. *.. @... . . . . . . . . . . . . . . ....................................
....................................

60 - l .. *..@ . . . . . . . . . . . . . . . .........
. . ...........
............. ..
..
.
.
.
.
.
.
trapsand escape to die later;the skin
..........
...........
..........
....................................
.. . . ............. . ??...........
?? ???
.... .... ....
...................................??
....................................?
....................................
.... .... ....
??? ??? ....
????? may be damagedeither in the course
..............................................
.................
.............
... .... . .. . .. .. . .. . .. . .... ... .. ... of the hunt or in processingor stor-
............
.... .... . .. . .. . ..??
....................................
....................................
.................................... .??? .. ?? . age; the animalmay be too small and
::::::::::::::::::::::::::::::::::::::::::::::
...................................? ::::::::::.::::.:::::
???? ??
50 - . . ...........
.
. ... ... . . . . . . .. ...........
...................................
...... ......
... ... .........
. .. . . ... .... ......
. . . . . . . ...........
... ... ......
..:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:-:.:.:.:.:.:.:.:.:.:.:.:-:.:.:-:.:.:-:
. .. . .. . ..
...... . .... -....-....-is discarded either by the hunter or
.....................................
. .... . . . . . . . . . . . .
.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-:-.-.-:
....................................
... *................................
. .
the buyer; or a lactatingfemale may
...................................???
....................................
....................................~? be killedand her infantdies as a result
Psittacidae(6/18) ..................................
...................................
....................................
....................................
.?? ?????
.. ??? ....??
6.2% ...................................
...............::::::::::::::::::::
...........
.:.................. .....................
....................................??
........ .. .........
....................? ?...?????~?
??
...
.... of her death. In the case of the live
. . . . . . . . . . . . . . . . . . . . . . . . .?
. ...................................
...................................
....................................?
..............................
...................................?
:::-::::::::::::::::W:::w:::-:::-:::-:::w::::::::::::::::::::
.................................... I ?. ?. ??
??. ???? . .?
?????. .I . . ...............
ww.
... .....**.v@.@@.**.@
animal trade, the mortalityin trans-
40 - Psophidae(1) ....................................
..................
....................................
. . . ...... . . ... . .. . ...
................................... .. . ......................
. .. ... . .. . ... .. . . ... . . . . ... . .........
....................................
....................................
.................................... ?? ? ?? ?~?~~?~ port to the point of final sale is even
4.7% ....................................
.................. .. .. ... .. ..

Phasianidae (1)
........ . ..
...................................
....................................
.. .......
. .. . . ..
*.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.-.:.:.:.:.:.:.:.:.:.:.:.-:.
.....................@
9.
. ..
??????? . .
??I ??? higher (Grimwood 1968).
....................................
...................................
....................................
........
.......................... ........ ........ ? ?? ?. .......
???
. ..... To illustrate the number of animals
5.2% ....................................
....................................
::-::-::-::-::-::-::-::-::-::-::-::-::-::-:-:-:-:-:::::::::::::::::::::
............................. ......?
....................................
....................................
....................................
....................................
? ? ?I? ~?? involved in the commercial wildlife
30 - ...................................
...............:::::::::::::::::::::::::::
....................................
....................................
....................................@
....................................?I ? ??~??? ??? trade, Table 1 lists the number ex-
Cracidae(4) ...................................?
....................................? ?? ????? ? between 1962 and 1967 from
...................................?
::::::::::::::::::::::::::::::::@:::::::::::::::::::::::
....................................? ? ??
?..........
?? ???~ ported
12.6% ...........
....... ................. ...... ...... ...... the Amazonianport of Iquitos.Using
................................... ???? ?
.:.:.:.:.:.:.:.:,:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:,:.:. the correction factor of three, the
20 - total numberof animalskilled would
be almost5 million,or approximately
800,000 animalsper year or one ani-
Tinamidae
(9) ---------I......................... mal for each square kilometerof the
10 - 18.8% PeruvianAmazon each year.
Game birds (29)
9.1%
Table 1. Animals and animal skins exported
fromIquitos,Peru,1962-1967. (FromRedford
0 and Robinson1991.)
Biomass NumberSpecies Numberof
individuals
(190kg/km 2) (319 total) Animals exported
Figure4. Avian diversityin AmazonianPeru.Numbersin parenthesesare numbersof Live monkeys 183,664
speciesin taxon.(DatafromTerborgh
et al. 1990.) Skins
Caiman
III Melanosuchus 47,616
killedan estimated10,000 smallspot- America (principallyArgentina,Bra- Caiman 101,641
ted cats in a 15-year period (Paz y zil, and Venezuela)exported 15,000 Mammals
Mifio C. 1988). kg of egret and heron feathers,repre- Capybara(Hydrochaeris) 67,575
FEATHERS.Much less well-studied Otter (Lutra) 47,851
senting an estimated 12-15 million Giantotter (Pteronura) 2529
is the exploitation of birds for their individualsof smallerspeciesof birds
Ocelot (Felispardalis) 61,499
feathers. Although currently out of and 3-4.5 million largerones. 9565
Margay(Feliswiedii)
fashion,featherswere once important BODY COUNTS. In interpretingthe
Jaguar(Panthera) 5345
elements in women's fashion. The numbers of individual animals in- Collaredpeccary(Tayassu
tradeconcentratedon egretsand her- volved in the commercialtrade ema- tajacu) 690,210
ons. At the height of the feather nating from the Amazon, previous White-lippedpeccary
frenzy,premiumfeatherswent for $5 authors used either the number re- (Tayassupecari) 239,472
Deer (Mazama) 169,775
per plume or $28 per ounce in New portedlyexported at various ports of Total
York.Between1899 and 1920, South exit or the numberimportedat vari- 1,626,751

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 It is importantto realize that sub- 100
sistence and commercialhunting oc-
:lill:ml-:llelil.l.l.'l......I.
b-11'llllll1111111lll
o. o%o.
..,.,...,,......... ,.......
. . .
.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:-:.:.:.:.:.:.:.:.:.:.:.:.:.:.:l
.oo~.o.. oo.o.. o.o. ooo.~OoO.oOO.o....o...
,.... ,... ,.....,..,..,.,...,
. . .
....
*i!i!i
i!!iii!iiii!i
iiii!i *
....................................

[!iii!!iii
... ......

....................................
............... .....................
... ... -.-.- ... -...-.-.-. -. --..
....

..............
....
o..o.o.oooo...
.... . . . . o .o o. oo. . . ... . . . . . ..o.o..o...
. . .
cur simultaneously.As a means of ....................................................................
..,.-.,...,
..............
.-.......-.,...........,......,...-.......-
........................ ...................
..........-...... Rrl utd.-.-.....-..- ................. .................... ...............

providinga firstapproximationof the .o.% ..o. . .

. .,.....,
....................................?:
..........-.
......-................,...,.-..,,.,......
. .... .
,..,....,..,.
.o .....o
.-.....,...,.....,.. o....~o
,........,.
...... .............
...................................
. . . . . . . . . . . .. . . . . . . . . . . . . . . ..
......................... .
. . . . . . .......................................

90 - ...................................???????
....................................?
.O...o.......
,.-.-.-.-.-.-... . .oo.oo..oo....o~o
. . .. . o.o.o.....
. . . .....ooo..oo.
......... ................................... ............................l
:-:.:.-::::::::::-:.::-- :-.. :.-- - :.;: -;..-:.-.--:.:.:.:.-.:-:
combinedeffectsof these two typesof ...................................???????????????
...................................
...................
......................................
.........................-.-.-.-.-.-.-.-..-
...................................
.-:.:.:.:-:.:.:-:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:-:.:.:-:.:.:.:.:.:.:.:.::-
................................... . .. . . . . . ......-............---------.
...................................
....................................
...
... .... ... ... .... ... ... .... ... ... ....
...-... -..-...-.-.-.-.-.-.-.-.-.-.-.-.-.-.............
..o..o-...-...... .. .. ....
... . . . . . . ...

......................................
o..o.-....-..
....
. ... .... .. . ... ... ... .... .. . .. ... ...
......
..... ............................

hunting, at least in the 1960s and

..........
. . . . . . .. . . . . . .......--------- ... ... ... ... ... .. . ... ... ... ... ... .. ............................
..........................................
..................
. . . . . . . . . ..........................................
. . . . . . .
... . . . .. . . .. . . .. . .
.....................................~ .......................................
....................................

1970s, I have extrapolatedthe above ...................................???

..........
. . . . . ............. .
..............................
,...................................
. .................................
...........
............................ ..

estimate to the 3,581,180 km2 Brazil- ...................................????????????

....... .. ...... .... .. ... ..... ......... ...
... ... ...
..
80 - ...................................??????????
.,.......... .......................... . . . . . . . . . ..........................................
. . . . . . .
....................................?? ..........
... .. ?
...........-.-.-.-.-.-.-.-.-..-
...... ?o... .o .. .-o.o. ??-. ??o.-.o??..........
??.- ??.- ?--
ian Amazonian states. This calcula- ....................................?? ..........
..... .. . ..
.........?????
.. . .. .. .

...................................???????? ...................................
..............-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-.-..-
.........l

tion yields an estimate of 4 million .

.........
.......
.
.. . . .. . . . .. . . . .. . . .
. .
.....
.
.............
..
... .-.-..
.. .. . . - .
--
. -
..
.
e
-
.
. ..-
..
..
.. .??
.
.

animals killed for commercial pur- .... ... .... ... ... .... ... ... .......... .........................
............ .... .... .... ....
....................................
.....

....................................
....................................

poses per year, which may be com-

.................-.-.-.-.-.-.-.-.-.-.-.-.-...

70 - ........
........
... ..... .. ........ ... .........
..... ... .. . .. . . .. ... ... ... ... .... .. ...................-..
......
... .. .. ..... .... ... .. .....
....................................

.... ... ....

. . . ... .. ...
....................................
. .. . .. . .. . .... .. ... .... ................................

bined with an estimated 19 million LargePrimates

(4)
....................................
...............
....................................
..................
...............-.-.-.-.-.-.

......................... ..............

killed each year for subsistence. 35.6% *e..-.. -.. -.-... .. ............
........ .. ... .. ... .. ... ..
1..7. 9?.
Therefore, approximately 23 million
!i!i!*!i!i!i!i!i!e!e!i!i!i!i!*!i!i!*!i!ii*!iii
. .. .. . ..
.. . ......... ......
.. . . .....
.......
.....
...
.. . ....
.. .
... ... . .. .. .. ... ....... ....................................

Large Primates (4 .. .. .. .. . .. .. .. . .. .. .. .
.................. ......................-----

animals are killed per year in the 60 - Alouatta=1 11.8%

Brazilian Amazonian states. If the Ateles=l
1.5% ...............

........ ........ ....

...
..... ... ........
.....................-.----

..
.... .. ......
correction factor for fatal wounding Cebus(2)=12.3% ...................................
.................
...
.. ....... ...
.............. .. .... ...
.. .... ... .. ............
. . . . .

....
. .
---------..........-
.

.... ....
.

is applied to subsistence hunting as ................... . . . . . . . .

50 -
...................................
....
....... ... .. . ... .. . .. .............
... .. ...... ....... ........ ....
...... ........ .. .
well, the total reaches 60 million.
:.:.--:.:.:.:.:.:.:.:.;:.:.:.-:.::-:.:.:-.:.:.2:.:.:.:.:-:.:.:-:-:.:-:-
.-:.:.:.:.:.:.:.:.:.:.:.:.-:.:.:.:.:.:.:.:-:-.:.:.:.:.2.:.:-;.:.:--.:-
....................................
... .......
...................................
. ........... .

Given the increase in human popula- . ....

..... ...--.-.
.. .......
...
...................................
-
..... . ....
.-.-.-.-.-.-.-
........ ...
tion in the Brazilian Amazon since ............ :.:.:.2
..
. . . . . .
.......... ...:.:n-am.:.:.:.:.:.:.:.:.:....:.-............
. . . . .
... ... . .. ............

1980, this total undoubtedly is an 40 -

..................................
......... ....... ......... ... .. .........
....
underestimate of current kill rates.
.....................
...... ...... ...... ...... .... ......
.................
LargeRodents(4) Game:--:: .................
Game... mammals
... ....
mammals
...................................
.....
..... .....
............. .............
....
:..::.:::.
........(2
.... ...
.... .....
....
.....
..... .....
...... .....
11.3% ...... ......
.... .... ......
...................................
.... .... .... .... ......
....
Which animals are killed
...... ... ... ... ... ... ... ...
by hunters? 30 - ..... .. . .. . . .. . . .. . .

Before attempting to assess the im- Perissodactyla(1)

7.3%
pact of the removal of such large
numbers of animals from Amazonian 20 -
forests, it is necessary to consider
several factors. First, the most com- Artiodactyla(3)
monly taken game animals are almost 21.0%
always the largest members of their 10 -
group and usually the largest species
in the forest. Hunters prefer birds and
mammals of large body size. The only U
large mammals not commonly hunted Biomass Number Non-Volant Species
for food (felids and otters) are hunted
for their pelts. Large wading birds (1526.6 kg/km2) (67 total)
and raptors are the only large birds
not commonly hunted for food. Figure5. Nonvolantmammaliandiversityin AmazonianPeru.Numbersin parentheses
are numbersof species in taxon. (Data from Terborghet al. 1986 and Janson and
Second, large animals, although Emmons1990.)
represented by relatively few species,
are major contributors to the overall
biomass. The best data available to 12 species make up 75% of the mam- tions of moderate hunting, densities
illustrate this point comes from the malian biomass (Figure 5). To show of nonprimatemammaliangame spe-
work done at Cocha Cashu Biological that the Manu site is not atypical of cies decreased80.7% when compared
Station in Manu National Park, Peru neotropical forest sites, Figure 6 com- with similar, unhunted sites. Under
(anson and Emmons 1990, Terborgh pares mammalian biomass data from hunting conditions describedby the
et al. 1990). Of the 319 bird species three other nonhunted sites. Clearly, authors as heavy, nonprimatemam-
recorded at this site, 9% are com- species that are preferred game make mal densities decreased93.7% com-
monly hunted (Redford in press, Red- up a large proportion of the biomass pared with similar, unhunted sites
ford and Robinson 1987). However, in unhunted sites. (for details, see Redfordin press).
these 29 species make up 52% of the Third, and as a result, if hunting Due to the greateramount of data
total avian biomass (Figure 4). affects the abundance of game spe- on primates,a similarcomparisonof
This pattern is even more striking cies, areas that have been hunted the effect of huntingon primatescan
for the mammals, for which data are should show decreases in density and include both density and biomass.
available only for nonvolant species. therefore biomass of these species. A Data from many differentAmazonian
Of the 67 total species, 18% of the review of the available data of effects sites show that in hunted areas large
species are commonly hunted. These of hunting shows that under condi- primate biomass drops 93.5% when

417
June 1992

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 1990) shows that 69.5% of the game
100- bird biomass is composed of fruit-
eating species. This total would in-
creaseto 96.2% with the inclusionof
non-hunted the curassow (Mitu),which some ex-
75 - perts believe is also primarilyfrugiv-
orous.1
Many of the most importantcom-
Edentata mercial food fish in the tropical for-
50 - est are also large fruit-eatingspecies.
Large Primates Many of these frugivores are in-
volved in seed dispersal,seed preda-
tion, and the structuringof tropical
LargeRodents forests.
25 -

Perissodactyla
The ecology of game animals
. Artiodactyla and of their absence
. . . . . . . . . . i......
. . . ..
0 __?__ _.
Manu, Peru Guatopo,Venezuela HatoMasaguaral, BarroColorado, Althoughmany ecologistshave docu-
Terborghet al. 1986 Eisenberg1980 Venezuela Panama mentedthe importantroles playedby
Eisenberg1980 Eisenberg1980
large animals in seed dispersal,seed
predation,herbivory,pollination,and
Figure 6. Biomass of game mammals as a proportion of total mammal biomass at four
neotropical sites. Only the animals indicated in black are not hunted.
predation, until recently few have
consideredthe role of largeanimalsin
tropicalforests and what would hap-
pen if they were removed from the
-

comparedwith similar unhunted ar- 50.9% to 26.8% of the total avian system (Emmons 1989, Janson and
eas, and large primate density drops biomass. Emmons 1990, Terborgh 1988).
80.7% (Table2). In fact, Freeseet al. A fourth factor that must be con- Some ecologists have stated that re-
(1982) have statedthat "predationby sideredbefore assessingthe effectsof moval of some individual species
humans is clearly one of the most the removalof large numbersof ani- from an ecosystem would probably
important factors affecting monkey mals from Amazonianforests is that not have any substantialeffectson the
densitiesin most of the Peruvianand in tropicalforestsmany of the largest remaining species (c.f. Feinsinger
BolivianAmazon, and probablyelse- animals,both terrestrialand arboreal, 1983). Some studies, however, have
where in South America"(p. 82). are frugivores.Turning again to the not only documentedthe existenceof
A similar comparison made for Cocha Cashudata, 84% of the game- keystone species but also have dem-
game birdsshows a drop of 73.5% of mammal biomass is composed of onstrated what happens when such
originaldensityundermoderatehunt- fruit-eating species, according to a species are extirpated. Recently,
ing and of 94.6% underheavy hunt- calculationtaking the biomass of the Brownand Heske (1990) have shown
ing. Similarconclusionswere reached most commonly hunted mammalian that the removalof a guild of kanga-
by Terborghet al. (1990) when com- taxa and considering frugivore- roo rats from an experimentalplot in
paring Panamanian and Peruvian omnivores,frugivore-granivores, and the southwestern United States re-
avian biomass. Finally, Thiollay frugivore-herbivores as frugivores sultedin majorchangesin the vegeta-
(1986) showed that in comparing a (data from Robinson and Redford tion structure.
hunted forest site with an unhunted 1986). Suchclear-cutcases are not known
forest site in French Guiana, avian A similaranalysisof the gamebirds from neotropicalareas, but there is a
game speciesbiomassdecreasedfrom (food habit data from Terborghet al. growingbody of work suggestingthat
in this ecosystem large vertebrates
Table 2. Impact of hunting on large primates in Amazonia. Means (standard deviations) are given. may perform important ecological
roles and that theirabsencewill result
Primate biomass* Large primate percentage of
a (kg/km2) total primate biomass* Number of sites in a changed forest (Janzen 1988).
Unhunted 9
Such a conclusion has been reached
363.8(314.3) 64.5(11.4)
Hunted 23.8 (38.1) 19.6(29.6) 19 for paleotropicalsettingsand in com-
parisonsbetween the Pleistoceneand
Primate densityt Large primate percentage of the present,but it has not been made
b (individuals/km2) total primate densityt Number of sites explicit for neotropicalsettings.Stud-
Unhunted 34.1 (10.8) 33.8(16.3) 7 ies that shed light on the ecological
Hunted 6.6 (5.8) 8.1(10.0) 8 functions of large neotropicalverte-
*Data from Peres 1990, Freese et al. 1982, and C. Mitchell and E. Raez Luna (unpublished results,
1991, Wildlife Conservation International).
tData from Peres 1990 and C. Mitchell and E. Raez Luna (unpublished results, 1991, Wildlife 'A. Grajal, 1991, personal communication.
Conservation International). Wildlife Conservation International.

418 BioScienceVol. 42 No. 6

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brates fall into three categories:her- in Panama,Howe (1984) stated that rainfallis due to the additionof bird-
bivory and seed predation, seed dis- "animal-mediateddispersalis certain dispersedand mammal-dispersed tree
persal, and predation. to be critical for the demographic species. In addition, he shows that
recruitmentof many or most tropical wind-dispersed species tend to be
Herbivory and seed predation. In forest species"(p. 266). wide-ranging, whereas mammal-
Mexico, Dirzo and Miranda (1990) Large birds, particularlythe tou- dispersedspecies tend to be localized
compared two tropical forests, one cans and cracids,are amongthe most in distribution.For example,43% of
with its full complement of large important seed dispersers.Many of the tree speciesfound at Rio Palenque
mammals(peccaries,deer, and tapir) the speciesof cracids,particularlythe were presumablydispersedby large,
and another in which these species curassows, are among the species nonvolant mammals, and 50% of
had beenextirpatedby hunters.There whose local populations are most these are endemicto coastal Ecuador
were strikingdifferencesbetween the rapidlydepletedby hunting.They are or to Ecuador and adjacentColom-
two forests. The hunted forest was also slow to reproduce,with the av- bia, whereas only one mammal-
typified by seedling carpets, piles of erage cracid requiring at least six dispersed species ranges throughout
uneatenrotting fruits and seeds, and years to replace itself in the popula- tropical America. Given these facts,
herbs and seedlings undamaged by tion (Silvaand Strahl1991). Because the loss of largevertebratedispersers,
mammalian herbivores-phenomena of the cracids' importance as seed which has undoubtedly occurred in
much less evident in the unhunted dispersersand susceptibilityto hunt- much of this area, would affect not
forest. ing, Silva and Strahl have suggested only local populationsof trees, but in
A second example comes from that "humanimpact on the Cracidae many cases it might result in the ex-
work done comparingseed and seed- may have irreversiblelong-term ef- tinction of tree species with only lo-
ling predation, tree recruitment,and fects on the biology of neotropical calized distributions, resulting in a
rodent populations on Barro Colo- forest ecosystems"(p. 51). loss of diversityof plant speciesmuch
rado Island, Panama,with adjoining The other groupof importantlarge greaterthan expected.
mainlandareas (De Steven and Putz seed dispersersare the primates,par-
1984, Glanz 1990). On the island, ticularlythe woolly and spider mon- Predation.The role played by preda-
ocelots and other large mammalian keys. In a study in Surinam,spider tors in structuringcommunitieshas
predators are absent, with agouti monkeys were shown to disperse been well studiedin marineand inter-
(Dasyprocta) and squirrel popula- seeds 93.5% of the times they fed on tidal systems. This work has shown
tions high comparedwith the main- fruit and apparently served as the that predatorscan increasethe overall
land, where not only felids, but also only dispersal agent for several tree species diversity in a communityby
humans,have reducedthe population species(von Roosmalen1985). Defler decreasingthe abundanceof smaller
of these rodents. Predationof seeds (1989) has shown a similar pattern predators and competing herbivores
and seedlingsof severalcanopy trees for woolly monkeys.Like curassows, and by reducingdominanceof plant
on Barro Colorado Island is much spiderand woolly monkeysare highly prey species.
higher than on the mainland due to prized game animals and are rapidly Such research has not been con-
high populationsof seed-eatingmam- huntedout of a forest(Peres1990). In ducted in neotropicalforests, but bi-
malson the island.In an elegantstudy the absence of such large primates, ologists working in various locations
of small (1-4 ha) islands in an artifi- many species of plants may experi- have observed that a decrease in
cial lake in Panama,Putzet al. (1990) ence severely altered seed dispersal abundanceof large predatorymam-
showed that, in the absence of seed- patterns. mals is correlatedwith the increasein
eating mammals, trees with large Interestingly,agoutis in additionto abundance of medium-sized terres-
seeds had a distinct advantage over being majorseed predatorsalso serve trial mammals, particularlyagoutis
those with small seeds and came to as seed dispersalagents. It seems that (Glanz 1990, Janson and Emmons
dominate the small forest patches in at lower densitiesthey are important 1990). Absence of large predators
less than 75 years. dispersalagentsfor large-seededtrees such as jaguars, pumas, and ocelots
Otherimportantseed predatorsare (c.f. Forget 1991, Hallwachs 1986); also seems to result in more uneven
peccaries, deer, and tapirs (Bodmer in their absence, at least some trees densities of prey species (Emmons
1989); these species consume an would become locally extinct. 1987). Da Fonseca and Robinson
enormous number of seeds, particu- Another important group of seed (1990) suggestedthat the absenceof
larly those from palms. These ungu- dispersers is tropicalfish. Fish appar- ocelots in forest patcheswas the rea-
lates, especiallythe white-lippedpec- ently disperse seeds of at least nine son for large numbers of opossums
cary with its large group size, were plant families(Gouldinget al. 1988). and consequent lower small rodent
probably important elements in af- Many of the fish seed-dispersersare densities.
fecting forest composition and struc- important food fish and are heavily Intactcommunitiesof largecats are
ture but now are very rare. pursuedby both subsistenceand com- rare in neotropical forests. Even
mercialfishers. where there has been no hunting of
Seed dispersal. Many authors have Gentry (1983) has shown, in the theseanimalsfor theirskins,therehas
documented the important role neotropics,that areas of higher rain- usuallybeengamehuntingfocusedon
played by large birds and terrestrial fall have highertree diversityand that the speciesthat are primaryprey spe-
mammalsin the dispersalof the seeds much of the overall increasein plant cies of the large cats. Large raptors
of tropicalplants. Based on his work diversity accompanying this higher are likewise affectedas they eat pri-

June 1992 419

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 and mammals, an indication of the
ecological roles played by these spe-
cies is possible.These examplesmake
clear the importance of fruit-eating
vertebratesin structuringand main-
taining tropicalforests. What exactly
will happen as a result of the loss of
the game animals is not clear. Some
cases, as on Barro Colorado Island
and in the Mexican forests, provide
hints as to these consequences-the
resultingforest will be determinedby
a complicatedmix of more predation
on some species, less on others, and
the rarificationor extinction of still
others. What furthercomplicatesour
abilityto predictoutcomesis the fact
that humanshave been alteringtrop-
ical forests all along. What has
changedis the intensityand scale.
Ecological extinction. During the
height of the skin trade, many ani-
mals with valuableskins were killed.
Since the collapse of the skin trade,
tropicalforestpeopleshave continued
to hunt many of these animals be-
cause their meat is appreciated.The
result of this widescale human activ-
ity has been the reductionor extinc-
tion of local game populationsin vir-
tually all areas of Amazonia.
Conservationbiologists are by def-
inition concerned about extinction.
Yet, as Estes et al. (1989) point out,
there are several types of extinction:
global extinction, local extinction,
and ecological extinction. Ecological
extinctionis definedas "thereduction
of a species to such low abundance
Figure7. A group of Braziliansliving along the AmazonRiverdisplayan ocelot skin. that although it is still presentin the
Photo: Kent H. Redford. communityit no longer interactssig-
nificantly with other species" (p.
marily animals that are major game resulted in the elimination of adult 253). Althoughof tremendousimpor-
speciesfor humans. otters from more than 20,000 km of tance, conservationists ignore the
There has been much less work waterway per year. Both black widespreadnature of ecological ex-
done on the effects of large mamma- caimanand giant otters, the two larg- tinction in neotropicalforests, focus-
lian and reptilian predatorson neo- est carnivoresin neotropicalfreshwa- ing instead on demographicextinc-
tropical aquatic ecosystems. Yet this ter ecosystems,have also been elimi- tion (extinction of a population or
guild has probablybeen the one most nated from many areas.The effectsof deme) and calculations of minimal
heavily affected by human hunting. the loss of these two predators is viable populationsizes.
For example, an estimated3 million unknown, although Fittkau (1970, Even if jaguars, woolly monkeys,
caiman, river otters, and giant otters 1973 in Best 1984) has speculated or large curassows have not gone
were removedfrom from Brazil,Co- that caimanserve an importantfunc- extinct in the wild, their populations
lombia, and Peru between 1962 and tion in nutrienttransferfrom terres- may have been reduced to such an
1969.2 Grimwood (1968) calculated trial to aquatic ecosystems and that extent that they no longer perform
that, based on the home-range of their absenceis correlatedwith a de- their ecological functions. What is
river otters, the mid-1960s commer- creasein the diversityand biomassof needed is movement beyond the ge-
cial harvestingof river otters in Peru fishes. netically based concern with demo-
By examiningstudiesof herbivory, graphic size to a new emphasis on
seed predation, seed dispersal, and minimum ecologically operational
2K.H. Redford,1992, unpublishedresults. predation by large neotropical birds population size that incorporatesin-

420 BioScienceVol. 42 No. 6

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teractionsbetween plant and animal tors and insectivores, as well as a game does not even appear (Myers
species. threefold decreasein the biomass of 1988).
The animalsthat are the most pop- frugivores.
ulargamespecies,and the ones whose Conclusions
populationshave most likely become humans,hunting,
Integrating
ecologicallyextinct, include the most and conservation In tropical forests, large animals are
important predators, the large-seed importantnot only as food for people
dispersers,and the seed predatorsin Almost everyremainingpiece of neo- but also as integralecologicalcompo-
neotropical forests (anson and Em- tropical forest has been affected by nents of forestedecosystems.If these
mons 1990). These largeanimalspro- humans during pre-Columbian ecosystemsare to continueas forests,
vide what Terborgh (1988) has re- times, during the rubber-boomera, providingall of the financial,ecolog-
ferredto as a "stabilizingfunction." during the golden era of the skin ical and aesthetic benefits currently
Black caiman, jaguars,and harpy ea- trade, or more recentlyby gold min- desired, then animals must not be
gles maintainthe incrediblediversity ers, timber extractors,ranchers,and ignored.Many large animalshave al-
of tropical forests through indirect farmers.Even where no sign of hu- readygone ecologicallyextinctin vast
effects,"the propagationof perturba- man habitation is to be found, rub- areas of neotropical forest-areas
tions through one or more trophic ber trees show the unmistakable with large, towering tropical trees,
levels in an ecosystem,so that conse- signs of having once been tapped, lush ferns, and beautifulorchids.
quences are felt in organisms that piles of open Brazil nut shells show We mustnot let a forestfull of trees
may seem far removed,both ecologi- the telltale mark of a machete, and fool us into believingthat all is well.
cally and taxonomically, from the bones of caimankilled for their skins Many of these forests are "living
subjects of the perturbation" (Ter- whiten on the beaches. dead" (Janzen1988), and, although
borgh 1988, p. 402). Today, hunting is an integralpart satellitespassing overheadmay reas-
Humans are among the most se- of all forest-basedactivities,be it lum- suringlyregisterthem as forest, they
verely affectedspecies. The effects of bering,fishing,or medicinalplantcol- are emptyof much of the faunalrich-
huntingon large animalsare not just lection. For those living in the forest, ness valued by humans. An empty
of concernto those interestedin jag- hunting is an essentialcomponentof forest is a doomed forest.
uars and bird-watching.Hunting is a feeding a family; for those venturing
tremendouslyimportantsourceof nu- into the forest to collect forest prod-
tritionfor millionsof neotropicalfor- ucts, it is also a necessarysubsistence Acknowledgments
est-dwelling humans-a "subsidy activity. I would like to thankAllynStearman,
from nature" (c.f. Hecht et al. 1988) In virtuallyall areas of neotropical FrancisE. Putz, Marianne Schmink,
without which many other so-called forest,game animalpopulationshave and PeterFeinsingerfor theirvaluable
sustainableactivities, such as rubber alreadybeen affectedby humanhunt- comments on this paper. Carol
tapping, would not take place. As ing. This patterncontinuesas human Mitchell and ErnestoRaez Lunagra-
Peres (1990) has shown, in one year populationsgrow and forestexploita- ciously agreed to make available
and a half, one family of rubbertap- tion increases. The trend in recent some of their unpublisheddata. This
pers in the BrazilianAmazon killed yearshas beento increasethe amount is a contribution from the Program
more than 200 woolly monkeys, 100 of land allocated to multiple-usear- for Studiesin TropicalConservation,
spider monkeys, and 80 howlers. As eas-including Indian lands, Man Universityof Florida.
manyare beginningto realize,gameis and the Biosphere Programme re-
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