Вы находитесь на странице: 1из 11

The Cultural Keystone Concept: Insights from Ecological Anthropology

Author(s): Simon Platten and Thomas Henfrey


Source: Human Ecology, Vol. 37, No. 4 (Aug., 2009), pp. 491-500
Published by: Springer
Stable URL: http://www.jstor.org/stable/40343990 .
Accessed: 05/11/2014 09:25
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

Springer is collaborating with JSTOR to digitize, preserve and extend access to Human Ecology.

http://www.jstor.org

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500


DOI 10.1007/sl0745-009-9237-2

The CulturalKeystoneConcept: Insights


fromEcological Anthropology
SimonPlatten ThomasHenfrey

Publishedonline:28 May 2009


SpringerScience + BusinessMedia, LLC 2009

AbstractThe conceptof thekeystonespecieshas a long


itsvalidityremains
in ecologicalanalysis,although
history
have recently
researchers
controversial.
Anthropological
coinedthe termculturalkeystonespecies,but have not
fromexisting
differences
demonstrated
any significant
of culturallyimportantspecies. We define
treatments
as
cultural
accordingto theirsystemicfunction,
keystones
level
of
in
roles
essential
complexany
maintaining
having
itywithina social-ecologicalsystem.Examplesinclude
bitter
cassavaconsumption
amonglowlandSouthAmerican
groupssuchas theWapishanain Guyana,and commercial
of carrotsin RurukanVillage in Minahasa,
cultivation
Indonesia.Theseexamplesarebothessentialat one levelof
withinthe domesticand village
systemicreproduction:
cassava
in
the
case, and carrotswithinregional
economy
is
centred
each
While
markets.
upon a singlebiological
itselfis notthisspecies,buta
keystone
species,thecultural
severalmaterialand non-material
complexincorporating
systemelements.
species Cultural
keystone
KeywordsCultural
Social-ecological

Biocultural
diversity
significance

systemsEthnobiology

S. Platten
of Kent,
of Anthropology,
University
Department
UK
Canterbury,
e-mail:s.j.platten@kent.ac.uk
T. Henfrey
{M)
DurhamUniversity,
of Anthropology,
Department
Durham,UK
e-mail:t.w.henfrey@durham.ac.uk

The KeystoneMetaphorin Ecology


In a stonearch,thekeystone
is themostimportant
element,
Removeanyotherstone,
and is functionally
irreplaceable.
and theresultis a smallerarch,or one of different
shape.
Removethe keystone,and the capacityto forman arch
stonesone can make a
goes withit. Fromtheremaining
a
a
wall
or
number
of structurally
stack,
simpler
pile: any
forms.
intothe
thekeystonemetaphor
Paine (1969) introduced
of predation
theeffects
by
describing
ecologicalliterature,
of intertidal
thesea starPisasterochraceuson thestructure
Removalof the sea starleads to competitive
ecosystems.
loss of
exclusionamongitspreyspeciesand considerable
a
P.
thus
in
ochraceus
the
performs
ecosystem.
diversity
the
to
function
essential
maintaining
unique ecological
systemin a morecomplex(in thiscase, speciesrich)state
of Paine'sremovalexperi(Paine 1966, 1969). Replication
statusis notuniversal,
mentsshowedthesea star'skeystone
but manifestonly under certainecological conditions
are
(Menge et al 1994). Ecological keystonefunctions
to any particularspecies,but dependupon
not inherent
context.
While Paine's (1969) originaluse of the keystone
metaphorrestrictedit to top predators,others have
used it to referto species at othertrophic
subsequently
otherecologicalroles,includingprey,
levels or fulfilling
and herbivores
(Mills
hosts,habitatmodifiers
mutualists,
et al 1993; more recentexamplesincludeWillsonand
Halupka1995;Knappetal 1999;Milleretal 2000). Other
writershave extendedit beyond individualspecies to
processesinvolvingabioticas well as biotic ecosystem
elements(Bond 1994), guilds (Terborgh1986), microhabitats(Levey 1990), and multispecies
complexes(Daily
etal 1993).Debatepersists,
notablyin thejournalEcology
) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

492

and Society,betweenthose who employthe keystone species.As faras we areaware,thefirst


example
published
in thiswayderivesfroma
metaphor
conceptbroadly(De Leo and Levin 1997; Khanina1998; to extendthekeystone
ofironwood
importance
Vanclay1999; Higdon2002), and otherswho considerit studyoftheecologicalandcultural
of
Southwestern
Desert
in
the
Sonoran
trees
Davic
and
Fanelli
over-extended
2000,
tesota)
1999;
(Olneya
(Piraino
and considerthe NorthAmerica(Nabhan and Carr 1994): "Ironwoodis
2002, 2003). Some criticsgo further,
notonlya keystone
considered
unsoundto be
and
too
speciesin itsdeserthabitat,
empirically
keystone
concept vague
economiesof
crafts-based
to
the
a
cornerstone
but
also
et
Hulbert
useful(Mills al 1993;
1997).
of
communities
Mexican
and
Indian
ofecologicalkeystones Seri
Sonora,Mexico",
A return
to thenarrowdefinition
suggestedby Paine's researchremainstrueto theoriginal (St. Antoine1994: 70). This usage of thetermis purely
and is not accompaniedby any attempt
architectural
metaphorin which the keystoneaffects metaphorical,
of cultural
definition
a
towards
it
so
as
to
ensure
below
stones
interactions
keystones.
systematic
among
physical
cultural
the
of
a
formalisation
for
Precedents
an
it
would
be
thattheyaremutually
keystone
However,
supporting.
to identify
intowhatis no more conceptlie in variousattempts
errorto readtoo muchsignificance
speciesof par- whethersymbolic,utilitarian
thana fortuitous
coincidencewiththe spatialmetaphors ticularculturalimportance
at quantisalient.One attempt
as
economic
or
of
Modern
used to describetrophicrelations.
culturally
concepts
cultural
of
an
index
is
and dynamicssuggestthata broader ficationof this phenomenon
ecosystemstructure
in
various
basedon awarding
canplayan important
categories
role, significance
points
conceptual
conceptofkeystones
of which the cleareststatementis Holling's extended (Turner 1988). Adapted to increase its cross-cultural
thishas beenusedto helpprioritise
"All ecosystemsare controlledand relevance,
plantspecies
keystonehypothesis,
al
et
for
and
habitats
and
of
a
small
number
1990).
animal,
(Stoffle
protection
key plant,
organisedby
the landscapeat different Garibaldiand Turner(2004a) extendthe conceptof
abioticprocessesthatstructure
salience in defining"culturalkeystonespecies" as "the
scales"(Holling1992: 478).
salientspecies thatshape in a majorway the
The culturally
Identifying
keystonesin practiceis moredifficult.
in thefundamental
ofa people,as reflected
cultural
roles identified
identity
by Hollingcannotbe
key organisational
medicineand/or
reduced to functionalnon-redundancy(Schulze and rolesthesespecieshave in diet,materials,
have
workers
nor
Grimm
1995:
1994:
enthusiastically
Many
502;
practices".
9-10),
ecological
spiritual
Mooney
of
dominance(Power et al 1996). A more sophisticated and uncriticallyadoptedthisconceptas an indicator
et
al
Gelcich
et
al
culturalimportance
definition
of a keystonespeciesis one "whoseimpactson
2006;
2006;
(Byg
Stercho2006). Othersemphasiseitsvalue as a conceptual
its community
are large,and muchlargerthanwould be
expectedfromitsabundance"(PowerandMills 1995: 184). and practicaltool forlinkagebetweennaturaland social
An attemptto quantifythis effectas an index of com- sciencesapproachesin conservation
(Higgs2005; Ticktin
or CI (Power et al 1996) gives too et al 2006).
munityimportance
canbe useful
thata keystone
Whiletheinsight
effectsto
littleaccountof threshold
and othernon-linear
metaphor
a
valuable
is
formations
of
cultural
the
in
an
for
effective
basis
one,
study
keystones.
identifying
provide
on
several
criticism
bears
Turner's
definition
and
Garibaldi
its
with
the
functional
Combining
qualitativeaspect
redundancyrequirement(Schulze and Mooney 1994: grounds.It is not clear how,if at all, culturalkeystone
salientor
fromthosethataresimplyculturally
502) salvages both to produce a workablefunctional speciesdiffer
In
Davic
of an ecological keystonespecies (cf. Kotliar economically
definition
2004). addition,we
(cf.
important
a
2000), whichwe can thenextendto includeecosystem do not considerthemto have adequatelydemonstrated
to theecologicalkeystone
elementsof othertypes.We thus definean ecological claimof metaphorical
similarity
as anyfunctionally
component concept,upon whichtheyjustifytheiruse of the term
keystone
uniqueecosystem
andTurner
whose removalwould lead to a structural
note,Garibaldi
2004b).On a different
simplification (Garibaldi
indextoaid
a
use
of
and
with
Turner
its
abundance.
quantitative
(2004a)propose
disproportionate
an
of cultural
identification
keystone
species, approachwe
to use
flawed.
One
considerfundamentally
attempt
reported
thesecriteriafoundthemimpractical
CulturalKeystones
(The Snow Leopard
2007).
Conservancy
in whichtheir
Cristanchoand Vining(2004) avoid spuriousquantifiinhabitecosystems
Manyhumanpopulations
of the
treatment
in theiralternative,
that
can
thus
be
cation
role
is
of
a
and
independent
keystone
species,
ecological
as
criteria
seven
see
cultural
societies"
also
keystoneconcept,identifying
designated"keystone
(Meilleur1994;
defined
of
not
Kahn
In
recent
the
O'Neill
and
definitive, "culturally
2000).
Kay 1998;
years,
diagnostic,though
to keystone
theygiveno
species".Like Garibaldiand Turner,
keystone
concepthas beenappliedto humanformations
cultural
as to whatdistinguishes
describe the culturalimportanceof certainbiological clearindication
keystones
} Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

HumEcol (2009)37:491-500

493

frommorefamiliar
the main cash crop,and partof a culturalkeystoneat a
concepts.The suspicioninbothcases is
thatanynoveltyis moreterminological
thanconceptual.
different
level of systemorganisation:that of village
Ellen (2006) notesthe similarity
of theseapproaches integration
intoregionalmarkets.Its place in the market
of the term"ethnobiological domain distinguishesit in importantways fromthe
to his earlierintroduction
thatby virtueof Wapishanaexampleand othercultural"keystonespecies"
keystone
species",definedas "organisms
tohumansmaybecomeecologically
theirusefulness
crucial previouslyidentified
in the literature,
and demandsmore
of entireanthropogenic
to themaintenance
detailed
elaboration.
environments,
subsistence
systemsand ways of life"(Ellen 2006: 259).
Ellen'smajorexampleis use of thesago palm(Metroxylon
sagu) by Nuaulu in Seram, for whom its nutritional Cassava as a CulturalKeystonefortheWapishana
and range of otheruses have far-reaching
importance
consequencesfor land use and social relations(Ellen For GuyaneseWapishana,as formanyotherAmerindian
1978,2004a, b, 2006). Definingkeystonesin relationto groupsof lowlandtropicalSouthAmerica,bittercassava
the structure
and dynamicsof social-ecologicalsystems (Manihotesculenta)is centralto the dietand subsistence
withtheuse of theconceptin ecology,but strategy
is consistent
(Henfrey2002: 84-93). Not only is it the most
to a singlelevel of system important
in the sago case is restricted
crop,in termsof allocationof land,agricultural
Ellen's labour,and calorificcontribution,
its effects
on matter
and
However,like those it anticipates,
organisation.
to
be
information
flows
a
cultural
extend
far
assumes
this
material
contribuequivalent
keystone
beyond
approach
and use, and thecomplexof values
to a singlebiologicalspecies,whichwe believelimitsthe tion.Its consumption
cultural andpracticethatsurround
affectsubsistence
value of thekeystoneconceptforunderstanding
them,strongly
social organisation,
seasonaland daily activity
formations.
strategies,
of identity
and wellbeing.This
We assertthata culturalkeystoneis not a biological schedules,and perceptions
oftencentred
upon pervasive influenceon practicesto which it is only
speciesperse, buta complex.Although
relatedmakes the bittercassava complexthe
a particularspecies, a culturalkeystonecomplex also
indirectly
elementin Wapishanasocietyat thelevel
includesnumerousothersystemelements,bothmaterial majorstructuring
of
domestic
and
Platten
andfitsourdefinition
ofa cultural
andnon-material
2006).
reproduction,
(Platten2005; Henfrey
not
27
of
essential
for
in that
at
least
Brosi et al (2007) identify
survival,
Although strictly
species plants keystone.
rolecouldconceivably
in Pohnpei,Micronesia. itsnutritional
be replacedbyanyofa
involvedin canoe construction
as a cultural
ofthisactivity
Theircharacterisation
keystone numberof otherstarchycultivars,it is essentialto the
andreproduction
ofsocietyin itspresent
form.
practiceemphasises,in our assessment,how cultural maintenance
in
knowlAs
has
been
observed
a
number
of
other
lowland
combine
biological
species,
keystonecomplexes
Amazoniangroups,(e.g., Mentore1983-1984; Riviere
practice.
edge,andtechnical
of Wapishanamaterial
like ecological 1987; Heckler2004), manyfeatures
culturalkeystones,
We proposedefining
social
life
their
culture
and
reflect
the
to
structural
and
literal
unique processing
according
keystones,
keystones
roles,withinsocial systems.We thusdefineculturalkey- demandsand culinarypropertiesof bittercassava. The
toolkitinvolvedin cassavaprocessing
stones as systemelementswith crucial non-redundant extensive
comprises
level of structural the most importantmaterialpossessions of a typical
in maintaining
functions
any particular
The followingsectionselaborateupon this Wapishanafamily.The constanttasksof peeling,grating,
complexity.
datafromourown squeezing and cooking cassava roots are the major
to ethnographic
withreference
definition
of femaletimeallocationand also providean
of
researchon subsistence
practices Wapishanapeople in determinant
forsocial interaction
forum
and
2000
1999
Southern
amongwomenand
important
Guyanaconductedduring1998,
and
transmission
of
in theMinahasan vertical oblique
knowledge.
2002), and marketcultivation
(Henfrey
how
NorthSulawesi,during2000,2001 and
Unique qualitiesof cassava productsdetermine
villageofRurukan,
2002 (Platten2005).
Wapishanamen provisionthemselvesforextendedstays
A bag
andfishing
a cultural
we identify
In eachofthesesocieties,
expeditions.
keystone awayfromhomeon hunting
- a coarsecassava meal- and jar of casareep,a
based arounda single species of cultivatedplant. For of farine
madebyboilingdowncassavawaterand
condiment
cassava treacly
anduse ofbitter
people,thecultivation
Wapishana
- are the only foods
flavour
to
used
and
household
is centralto the subsistenceeconomy
soups and stews
with
into broadersocial networks.Its structuring normally
carried,supplemented hotpeppersgrownat
integration
over
are diverse;its importance hunting
effectson social relationships
camps,andeatenwithgameandfishprocured
to thediet, thecourseof theexpedition.
contribution
thusextendsfarbeyonditsmaterial
Cassava also playsa keysocialroleas thebasis forthe
which exemplifiesthe systemicpropertieswe treatas
alcoholicdrinks,notablyparakari,a thickbeer
of culturalkeystones.In Rurukan,carrotsare preferred
definitive
) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

494

in lower
Cultivation
without
meansofirrigation.
cassava roots.Parakarior households
made frommashed,fermented
at higher
othercassava-baseddrinksare crucialnotonlyto a good elevationfieldsthussupports
production
vegetable
altitudes.
which
in
collective
work
also
but
upon
arrangements
party,
achieve
most people depend for accomplishinglabour-intensive AmongRurukanvillagers,carrotsconsistently
tasks. One extendedfamilywhose religiousaffiliation high scores in freelistingexercises,and are the default
if no particular
about agriculture:
is notonlyexcludedfromthese topic of conversations
obligesthemto be teetotal,
assume
is
others
workgroupsbutalso sociallyisolated,
theyare
generally
commenting crop specified,participants
a
reflects
salience
cultural
This
them.
high
that,"theydon'tdrinkparakarilikeus".
talkingabout
bittercassava is thus keystonerole thatderivesfrom,but greatlytranscends,
substirutable,
Althoughmaterially
intotheWapishana.
Duringdrought- theireconomic importance.Collectiveorganisationfor
culturally
irreplaceable
in Rurukan,alongwithvillagespecialinduced cassava shortagesin 1998, even people with carrotproduction
in
lack
isation
the
of
other
foods
general,can be understoodas expressions
regarded
starchy
adequatesupplies
of the
Whilea Wapishanafamily of an egalitarianculturalmodel characteristic
of cassavaas a severehardship.
value
on
that
heartland
it
Minahasan
and
its
in
subsist
without
cassava
cooperation
places
could, theory,
products,
or and reciprocalaid (Platten2005, 2007). A Minahasan
would by doing so finditselfsocially marginalised
excludedin severalrespects,
as, "A personlivesto give
manyofwhichhaveimportant phrasewhichmaybe translated
of an
economicfunctions.
These includecollectiveagricultural lifeto others",accurately
expressesthecombination
and domesticlabour,communaleatingon huntingand emphasisupon sharingand the celebrationof individual
and manyothersnot describedhere advancementthatis at the heartof the region'smoral
fishingexpeditions,
such as germplasmexchangeand celebratory
drinking economy.
vehicleforexpresis an appropriate
such
Carrotcultivation
influences,
affecting
partiesor feasts.Thesestructural
well to the
and
conforms
this
sion
of
a rangeofsocialandcultural
of
subsistence
model,
particularly
beyond
aspects
harvest
A
Minahasanself-image. single
itsdirectmaterial
makethecultivation
anduse
generatesonlya
importance,
modestincome:the real financialbenefitsaccrue from
ofbitter
cassavaa cultural
keystone.
multipleannualharvests,requiringcooperationamonga
of specialisedworkersincludingcultivators,
community
truckdriversand markettraders.
Carrotsas a CulturalKeystonein Rurukan,Minahasa
wholesalers,
harvesters,
Labour is organisedthroughmapalus- reciprocaland
- in orderto preparethe ground,
In highlandareas of the Minahasa region of North waged work groups
raisedbeds,weed,harvest,
Sulawesi,Indonesia,the predominant
process,andtransport
activityis market construct
of vegetables.Mostvillagesoccupya particular thecarrotsfromthefields.
gardening
carrotproduction
marketniche,specialisingin the production
of a single
providesmanyopporConsequently,
in the village of Rurukan tunitiesfor collective labour, which along with role
crop.In the 1980s, cultivators
withinthe village.
cultivation
of carrots specialisationfostersinterdependence
beganto specialisein thecommercial
and market
moral
between
at
this
interface
Rurukan
now
the
sale
and
Success
(Daucus carota).
dependsupon
cultivation
of carrotsforbothliteraland symbolicpartic- economiespromotesindividualwealthand at the same
withcreatingstronginterpersonal
of commercial
ipationin regionalnetworks
exchangeand, timeis commensurate
anddirectsabotage
fortheassertion
of identity
and statuswithin bonds.Villagersemploygossip,rumour
by extension,
while
theregionalmarketsystem.
profits
hoarding
againstthosesuspectedof selfishly
andwealth
to theconditions
on theircontributions
Plotsurveysin Rurukanfieldsindicated
carrots
to be the skimping
as a whole.Thesetacticssimultaneously
dominantcrop. Several othervegetablesare extremely of thecommunity
collective
andpromote
innovation
andconformity,
onions
maize
common,including
{Zea mays),bunching
encourage
the
within
and
various
brassicas
action
{Allium
(Brassicachinesis,
village.
fistulosum),
Brassicapekinensis,
Brassica oleracea),each of whichin
Carrotcultivationalso plays a centralrole in the
someway supports
carrotcultivation.
The brassicassoften transmissionof culturalknowledge,both specific to
thesoil priorto plantingcarrots;bunchingonionsservea
and of widerrelevance.Technicalknowledge
agriculture
of
includesthe construction
similarpurpose,or can be interplanted
withcarrots
andsold specificto carrotcultivation
incomebetweencarrot raisedbeds, and particular
regimesof sowing,thinning,
piecemeal,providinga subsidiary
harvests.
withcarrots,
theplot weeding, harvestingand processing.Collective work
Maize, intercropped
protects
fromdraft
animals.It is also plantedas a cashcropin lower groups are the communitiesof practicewithinwhich
elevation
fieldswherecarrots
arelessproductive,
alongwith membersof the youngergenerationlearn to become
which
as well as themainsourceofincomeforyoung
household
income
cloves,
supplement
throughout cultivators,
the year and are the sole dryseason incomesourcefor adultsand newlyformed
households.
) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

495

Carrotsalso playa centralrolein thecombinedharvest


festivaland foundinganniversarycelebrationsin the
church.The climaxof the festivities
village'sProtestant
on a tablebeforethepulpitduring
involvesplacingcarrots
unlike
an elaboratepujian (act of praise). Significantly,
to guardian
othercropssuchas rice,thereareno references
intothe harvest
spiritsof carrots.They are incorporated
festivalas symbolicof the agricultural
systemand the
rather
thanas an object
actionsofthecommunity,
collective
of religiousworshipin themselves.
to Rurukanidentity,
Whilecarrotsaretherefore
integral
Thatis
for
thatidentity.
than
substitute
rather
theyrepresent
to say, while carrotsthemselvesdo not appear in any
andsale aresocially
theircultivation
Rurukan
iconography,
symbolicvis-a-visothervillages.Theyprovidethevillage
not only witha source of income,but with culturally
andstatuswithintheMinahasaRegion.
identity
appropriate
Othervillages,fortheirpart,occupysimilarly
specialised
economic niches. They thus exercise their keystone
intothisregional
at thelevelofvillageintegration
function
system.
JawedPeccaries:CulturallySignificant,
Whitebut notPart of a KeystoneComplex
the
Anotherexamplefromthe Wapishanademonstrates
betweenculturalkeystonesand specieswhich,
difference
are not implicatedin any
important,
althoughculturally
keystonefunctions.The white-jawedpeccary,{Tayassu
pecari),whoseWapishananameis bich,is themostsalient
and popular game animal for Wapishana subsistence
on sightingsof bich herds and
hunters.Information
are themaintopic
their
of
discussion
possiblemovements
of Wapishana
In freelists
of conversation
amonghunters.
and inevitably
animalnames,bichwas almostalwaysfirst,
mentioned.
names
three
two
or
first
the
among
withhunters
Interviews
suggestedkillratesforbichto
be among the highestfor all huntedmammals,and
over any other
to huntthemtake priority
opportunities
of resource
and
task.Huntingstrategy symbolicregulation
use may,to some extent,be adaptedto thebehaviourof
peccaryherds,whichrangeovervastdistances.
white-jawed
fromthatof otheranimalsin
differs
treatment
Its symbolic
toexcess(Henfrey
than
rather
hunting
deterring,
encouraging,
2002: 101-108).
we do not considerTayassu
Despite its importance,
cultural
keystonecomplexamong
pecari to be partof any
the Wapishana.If it did not exist,therewould be no
andotheraspects
behaviour
effects
uponhunting
significant
concerned
of cultureand societybeyondthosespecifically
is farfromexclusive:
withitsuse. Its dietarycontribution
therearemanyotheravailablesourcesofprotein,
including

a wide range of game animals,fish,various gathered


animalssuchas tortoise(mainlyGeochelonedenticulata),
and domesticlivestock.
A type of naturalexperiment
arises fromits patchy
in space and time,whichmeans it may be
distribution
absentfromcertainareas of foreston a temporary
or
basis.
When
for
several
no
permanent
years white-jawed
peccarieswerefoundin thehunting
groundsof thestudy
village, no dramaticchanges ensued, althoughpeople
lamentedtheirabsence and chargedtheirmostpowerful
shamanswithbringingabouttheirreturn.In some other
on fishand
villages,whose residentsrelypredominantly
domesticlivestockforanimalproteinand wherehunting
is
most
consider
relatively
unimportant, people
white-lipped
peccaries not as prized game, but a dangerousand
voraciouscroppest.
Developingthe CulturalKeystonesConcept
Each of the foregoingexamplesin some way illustrates
differs
how ouruse of theconceptof theculturalkeystone
fromthatof earlierauthors(Ellen 2006; Cristanchoand
Vining2004; Garibaldiand Turner2004a, b), in three
relatedways. Aside fromEllen, theydo not adequately
betweenkeystonesas systemelementswith
distinguish
functional
and speciesimporwell-defined
characteristics,
tantin otherways. All referto culturalkeystonespecies,
whereasthe culturalkeystoneis in facta complex,often
based aroundone or morebiologicalspeciesbutincorpoelements,both materialand nonratingseveral further
material.All also relate largelyto subsistence-based,
economicallyboundedsystems,and do not addresskeystone functionsrelatingto participationin supralocal
of commercial
networks
exchange.
The Tayassupecari examplehelps to illustratehow
fromotherforms
differs
in culturalkeystones
involvement
For the Wapishana,bich would
of culturalimportance.
score highon all ratingsof culturalimportance.
Despite
has
in
its
decline
nor
its
neither
absence,
this,
exploitation
hunt
Failure
to
effects.
or
cultural
wider
social
any
peccariesand eat theirmeatwould appearfairlypeculiar,
not to use
but have no deeperconsequences.In contrast,
cassava wouldimplyexclusionfromalmostall aspectsof
in thesubsistence
sociallifeand effective
non-participation
economy.
and
BothGaribaldiand Turner(2004a) and Christancho
lists
of
characteristics
or
Vining(2004) proposetypologies,
that identifycandidatesfor culturalkeystonespecies.
cases
Neitherprovidesanyreliablemeansofdistinguishing
elementin a cultural
suchas cassava,whichis an important
frompeccary,whichis not.In theRurukancase,
keystone,
thecentreofa cultural
thesameis trueforcarrots,
keystone
^ Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

496

HumEcol (2009)37:491-500

complex, and rice. Althoughit scores highlyon all


measuresof culturalimportance,
ricehas no keystonerole
(Platten2005). No approachof thistype,based on listsof
can be usefullydiagnosticof culturalkeystones,
features,
whichin the absenceof a formalsystemsmodel can be
identified
to contingent
featuresof any
onlyin reference
case.
Just
as
particular
ecological importanceis not
to
an
equivalent being
ecological keystone(Hurlbert
is not sufficient
to identify
a
1997), culturalimportance
cultural
keystone
complex.
CulturalKeystonesas Complexes
A further
limitation
oftheuse of listsofdiagnostic
features
is thattheseapplyto singlebiologicalspecies.We believe
thisto perpetuate
a crucialerrorin previousapproaches,
an
thatcultural
willequatewithbiologassumption
keystones
ical species.Both of the examplespresentedhereare of
culturalkeystonesbased aroundsingleplantspecies.The
to use thisspeciesas a signifier
forthewider
temptation
is
a
complex compelling singlespeciesis, indeed,"good
- butimpliesa reductionism
to think"
thatundermines
the
of
the
utility
concept.
the culturalkeystoneconceptin systemic
Formulating
termsdemandsthatit refernot to biologicalspecies,but
to complexesof interconnected
materialand subjective
factors.The social and culturalimportance
of Manihot
esculentaforthe Wapishanadependon the knowledge,
andtoolsinvolvedin itscultivation
anduse,along
practices
withseveralwildorsemi-domesticated
speciesthatin some
in the Rurukanagricultural
way supportthese.Similarly,
systemthekeystoneis notDaucus carota,buta complex
based aroundits cultivationand sale. The lower field
maize-cloveassociation,mapalusworkgroups,systemof
landtenure,
local egalitarian
ofthe
ethics,andthecentrality
churchwithinthecommunity,
areall essentialto theroleof
the carrotcultivation
the cultural
complexin maintaining
and
with
which
it
is
consubstantial.
The
identity practice
of Rurukanas a community
with distinct
reproduction
roleand statuswithintheregionalsystemdepends
identity,
of thiscomplex,whichcarrots
uponthekeystonefunction
alonecouldnotfulfil.
Extendingthe culturalkeystonefroma conceptually
securebutanalytically
notionof equivalence
unproductive
witha biologicalspecies createsa superficialsimilarity
withJulianSteward'sculturalcore,"the constellation
of
featureswhich are most closely relatedto subsistence
activitiesand economicarrangements...
(including)...such
as are empirically
social, political,and religiouspatterns
determined
to be closely connectedwiththese arrangements"(Steward1955: 37). However,thisdiffers
crucially

fromthe culturalkeystonein its specificreferenceto


it
As a result,
to theecologicalsetting.
cultural
adaptations
embodiesa seriesofimplicit,
assumptions
unacknowledged
"culture"(Ellen 1982: 52-65). We,in contrast,
concerning
simplyassumecultureto be a featureof any systemin
which humans play a part. The exact definitionor
to the present
characterisation
of cultureis unimportant
a
We
have
defined
cultural
keystone
complexin
argument.
such
role
within
termsof its structural
or organisational
culturalor culture-bearing
systems,withoutpreconceived
as to ecologicalor otherfunction.
assumptions
The incorporation
of subjectivefactorsintothecultural
also
highlightsa more important
keystonesconcept
of
our
analysis.The keystonestatusof any
implication
is
neither
inherent
nor inevitable:it is culturally
species
in
Rurukan'sspecialisation
and
this
ascribed,
contingently.
recentdevelopment.
carrotcultivation
is a relatively
During
thevillage
thelate nineteenth
and earlytwentieth
century
was renowned
foritsproduction
of palmsugar;eitherside
the
of Japaneseoccupationin the mid-twentieth
century
in
maize.
Thus
was
known
for
the
of
its
village
quality
it
is
not
of
the
the
details
cultural
Rurukan,
keystone
- but its veryexis- carrotsbeing substitutable
complex
tenceto whichwe ascribeadaptivefunctionality.
Neighbouringvillageshave similarcomplexescentredon other
It
orevenon non-agricultural
economicactivities.
cultivars,
is notthebiologicalspeciesinvolvedthatis crucialto the
ofthevillagewithin
and economicsecurity
status,identity,
the wider regional system,but the culturalkeystone
complexof whichit is a part.
In both cases, organisingeconomicproduction
on a
a consciouschoice.This
keystone
appearsto reflect
pattern
is obviousin thecase ofRurukan,
in whichthedecisionto
was sufficiently
recentto
specialisein carrotproduction
allow its documentation.
The originsof the Wapishana
obsessionwithcassavaaremorehistorically
obscure.Howit
is
one
of
of
sources
ever, only
many
complexcarbohydrate
whichraisesthequestion
of
cultivators,
grownbyWapishana
whya singlefoodplanthas beenassignedsuchan exalted
role.Itseemsthatinthiscasetoo,an economically
important
ofa cultural
via
plantspecieshasbecomethecentre
keystone
a consciousprocessofsocialengineering.
This observationanticipatesideas withinecological
on the potentialrelevanceof the keystone
engineering
model to social change.The widespreadimportance
of
in ecologicalsystemsis testimony
to theeffeckeystones
tivenessofa keystone
as a strategy
in engineering
structure
design (Rosemundand Anderson2003). This strongly
suggeststhatkeystoneswill also be commonin social
formations
(Lawtonand Jones1995: 147). In ecological
ithasbeensuggested
thata focuson keystone
management,
be
an
effective
means
of articulating
between
speciesmay

) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

HumEcol (2009)37:491-500

497

species-focusedand systemor landscape approaches


(Simberloff1998). Attemptsto base conservationof
Amazonianforestsupon the commercialextractionof
Brazil nuts (Bertholletiaexcelsa) show how flagship
based arounda keystonemay assistthereconstrategies
ciliationof ecological,social, and economicaspects of
amount
conservation
resource
(Clay 1997). Such initiatives
sometimes
toself-consciously
keystone
choosinga cultural
- as a strategy
for
centredupon an ecologicalkeystone
action,and appear to replicatean approachchosen in
and amongthe Wapishanaor
Rurukanin livingmemory,
in timespriorto theirrecorded
theirancestors
history.
CulturalKeystonesand SystemStructure
The role of carrotsin a culturalkeystonecomplex in
Rurukanderivesnotfromsubsistence
use, but fromtheir
cultivar.
Cristancho
commercial
statusas themostimportant
and Vining(2004) give no examplesof culturalkeystone
in the marketdomain,nor criteria
speciesso entrenched
of
relatedto thedegreeof commercialisation
significantly
Such analyseswouldmostlylikely
thespeciesconcerned.
saleas a singleuse.However,
formarket
cultivation
consider
numerous
canbe utilisedtowards
remuneration
thefinancial
a formal
in
economic
are
of
which
all
not
ends,
immediately
commercial
the
circumstances
in
some
and
major
sense,
Thisis notalwaysthe
keystone.
activity
maybe thecultural
case: amongthe Wapishana,the only cash crop of any
is peanuts,whichmanypeoplegrowto
notableimportance
thiscrop,andtheincomeit
raisea modestincome.However,
is
of
comparedto subsecondaryimportance
provides,
to their
farmers
and
sistence
alwaysgivepriority
production,
weaknessof such
cassava farms.This reflectsa further
of a cultural
generalisinganalyses: the characteristics
to another.
fromone situation
strongly
maydiffer
keystone
boundedsocieties,
Analysesbiasedtowardsconceptually
intermsofsubsistence
self-sufficient
produceconomically
cultural
of
the
to
tend
keystones
tion,
ignore possibility
and the
operatingat the level of marketintegration,
Market
circumstances.
different
ofthesevery
consequences
networks
in widersocial
impliesinvolvement
production
In
beyondthegroupforwhichthekeystoneis identified.
as a cultural
functions
carrot
keystone
Rurukan,
production
and because of its
at the level of thesewidernetworks,
interactions
market
in
the
upon whichthese
importance
are based,beingessentialforcommunity
networks
reproductionwithinthecontextof a regionalsystemof market
anduse
cassavaconsumption
exchange.FortheWapishana,
exerts its keystonerole at a lower level of system
withina largely
thatof domesticreproduction
complexity,
village-basedsubsistenceeconomv.In both cases, the

culturalkeystonelimitstherangeof culturally
appropriate
formsof social action,and thussimplifies
social decision
makingon thepartof individual
producers.
Thebenefits
areclearestintheMinahasancase,inwhich
it is easy to imagine alternativeways of organising
involvementin the regional marketeconomy.Basing
marketproduction
arounda keystonecomplexnot only
reflects
cultural
values,buthas manystrategic
underlying
advantages.At a local level, specialisationallows for
collectiveorganisation
of labourand sales,avoidscompetitionwithothervillages,andpromotes
thedevelopment
of
a greaterdepthof task-specific
skills.At a regionallevel,
productionis diverse, comprisingnumerousdifferent
in each of whichtechnicalcapacity
villagespecialisations,
a
is higher than similar numberof more generalist
wouldprovide.
strategists
It is clearfromthesedescriptions
thatwe regardcultural
as conservative
elementsof social systems.The
keystones
stablesocialformations,
and fixed
advantagesofproviding
for
individual
are clear,
reference
points
decision-making,
but dependupon otherpartsof the systembeingflexible
Theircontribution
and responsiveto changingconditions.
to the necessarybalancebetweenrigourand imagination
(Bateson 1979: 191-197, 204-219), means thatcultural
enable such dynamismelsekeystonesby theirstability,
wherein the system.However,it is clear thatany such
dominantsystemelementcould prove malintransigent,
in
the
faceof rapidexternal
changeifit limitsthe
adaptive
overallcapacityto adapt.
This has happenedon severaloccasionsin the recent
historyof Rurukan,whichhas seen severalchangesin
the identityof the dominantmarketcrop. This has
necessitatedprofoundchange in systems of market
cultivationand sale: the culturalkeystonecomplexhas
had to be rebuilton every such occasion. The global
faces a similarsituationin the form
economycurrently
of "carbonlock-in"(Unruh2000)- a set of interlinked
social, technical and economic factorsthat remains
stubbornlyresistantto change, and greatlyhampers
attemptsto reduce global dependenceon fossil fuel
As the culturalkeystoneconceptdevelops
consumption.
it may well contributeto improved
in sophistication,
of
understanding such situations,and provide useful
towardstheirresolution.
suggestions
as an elementthatsupports
keystone
Defininga cultural
some level or otherof systemcomplexity
suggestssteps
The use of
towardsa morerobustoperationaldefinition.
for
a
is
models
technique the nonpromising
ecological
whose
of
identification
ecologicalkeystones,
experimental
withotherspecies in the ecosystemare
totalinteractions
strong(Tanneret al 1994). A similar
disproportionately
flows
approach,based on formalmodels of information

) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

498

withinsocialsystems,
maywellprovidea practicalmethod the value of the conceptas a tool for interdisciplinary
to identifyculturalkeystones,as well as more precise communication.
However,the absence in mostcases of
itto little
raisesthedangerofreducing
criticalengagement
of theirsystemic
characterisation
properties.
can
aroundwhichinteractions
morethana catchphrase,
A morethorough
neverbe morethansuperficial.
working
and limitations
out of themeaning,significance,
Conclusion:The Importanceof the CulturalKeystone
potential
the
cultural
of
hamper
keystoneconceptmay,ironically,
Concept
the verysimplicity
such communication,
by eliminating
In summary,
cultural
arecomplexesofbeliefand that,like thatwhicha culturalkeystonebringsto social
keystones
itand limitsitsdepth.Thiswill
bothfacilitates
functionscrucial to social integration,
practicewith non-redundant
be worthwhile
at anyorganisational
scale withina sociocul- ultimately
if, by providinga conceptof
reproduction
turalsystem.Althoughmost documentedexamplesare genuinelydistinctive
analyticalvalue,it encouragesmore
baseduponone or a smallnumberof biologicalspecies,it meaningfulcommunicationamong the diverse fields
is not technically
necessarythatthis be the case. Most relevantto the protectionof biological and cultural
and
a
cultural
leadingto a deepermutualunderstanding
keystoneis in no case directly diversity,
importantly,
engagement.
capacityforproductive
greater
equivalentto a biologicalspecies.Even ifbased arounda
singlespecies,a culturalkeystoneis a complex,whose
to systemstructure
contribution
also dependsupona range Acknowledgements Thanksto Roy Ellen and Michael Fischerfor
advice and commentsduringthe writingof thispaper,to
instructive
of otherfactors,includingotherbiologicalspecies,arte- Rachel Kaleta for
introducingus to the keystoneconcept,to our
it friendsin each of the researchlocations,and to two anonymous
and socialpractices.
Moreimportantly,
facts,knowledge,
also dependsupon a rangeof purelysubjectivefactors: reviewers.Simon Platten'sfieldworkwas supportedby an ESRCThomas Henfreywas supportedduringfieldwork
social identity NERC studentship.
beliefs,ideas,normsand valuesconcerning
fromEC DG VIII, and duringwritingby a
an APFT studentship
by
and itsenactment
through
culturally
appropriate
practices. HuntpostdoctoralfellowshipfromtheWennerGrenFoundation.
thecultural
Extending
keystones
conceptfroma species
to a complex leads to further
observations.First,the
of thespeciesinvolveddo not
inherent,
objectivefeatures
determine
thattheywillbe partofa cultural
such References
keystone:
roles are culturallyprescribed,althoughit is entirely
possiblethatsome species are bettersuitedto thisthan Bateson,G. (1979). Mind and Nature:A NecessaryUnity.Fontana,
London.
others. Second, the detailed featuresof the cultural
Bond, W. J.(1994). Keystonespecies.In Shultze,E. D., and Mooney,
thanthe factof its
keystonecomplexare less important
H. A. (eds.), Biodiversityand EcosystemFunction.Springer,
existence.A keystonearrangement
forsocial integration
is
Berlin,dd. 237-253.
a functionally
sound one, on whichnumerousdifferent Brosi,B. J.,Balick, M. J.,Wolkow,R., Lee, R., Kostka,M, Raynor,
W., Gallen, R., Raynor,A., Raynor,P., and Ling, D. L. (2007).
societies have converged.Our analysis suggests that
CulturalErosionand Biodiversity:Canoe-MakingKnowledgein
societieshave, in effect,"chosen"to organisearounda
Pohnpei,Micronesia.ConservationBiology 21(3): 875-879.
keystonemodel,and this has shapedthe way in which Byg, A., Vormisto,J., and Balslev, H. (2006). Using the Useful:
Ecuador. Enviof used Palms in South-Eastern
Characteristics
particular
economically
important
speciesareused and not
8: 495-506.
ronment,
Developmentand Sustainability
thereverse.
J. W. (1997). Brazil nuts: the use of a keystonespecies for
In bothexamplesof culturalkeystonesdescribedhere, Clay,conservation
and development.In Freese,C. H. (ed.), Harvesting
themajormaterialelementis potentially
In
substitutable.
Wild Species: Implicationsfor BiodiversityConservation.The
JohnsHopkinsUniversity
Rurukanand otherMinahasanvillages,marketspecialisaPress,London,pp. 246-282.
tion in a singlecrop is a mechanismforeconomicand Cristancho,S., and Vining,J. (2004). CulturallyDefined Keystone
Species. HumanEcology Review 11(2): 153-164.
ofthevillageintoa regionalsystemof
symbolicintegration
Daily, G. C, Ehrlich,P. R., and Haddad, N. M. (1993). Double
commercial
exchange(Platten2005, 2007), and a locus of
keystonebirdin a keystonespecies complex.Proceedingsof the
NationalAcademyof Sciences of the UnitedStatesof America
articulation
betweenindividualproducers
and thisbroader
90(2): 592-594.
and Platten2006). FortheWapishanathe
system(Henfrey
R. D. (2000). Ecological Dominantsvs. KeystoneSpecies: A
Davic,
dominanceof cassava as a staplefacilitates
social integraCall for Reason. ConservationEcology 4(1): r2. http://www.
tionat thelevelof thedomesticeconomy.
consecol.org/vol4/issl/resp2.
The majorimpactto dateof previoustreatments
of the Davic, R. D. (2002). Herbivoresas KeystonePredators.Conservation
Ecology 6(2): r8. http://www.consecol.org/vol6/iss2/resp8.
culturalkeystoneconcept has been their uptake by
R. D. (2003). Linking Keystone Species and Functional
Davic,
in conservation
researchers
biology(e.g., Byg et al 2006;
Groups:A New OperationalDefinitionof the KeystoneSpecies
Brosiet al 2007; Gelcichet al 2006; The Snow Leopard
Concept. ConservationEcology 7(1): rll. http://www.consecol.
2007; Ticktinet al 2006). Thisdemonstrates
org/vol7/issl/respll.
Conservancy
) Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

499

Khanina, L. (1998). DeterminingKeystone Species. Conservation


Davic, R. D. (2004). Epistemology,Culture,and KeystoneSpecies.
Ecology and Society9(3): rl. http://www.ecologyandsociety.org/ Ecology 2(2): R2. http://www.consecol.org/Journal/vol2/iss2/
resp2.
vol9/iss3/respl/.
D.
De Leo, G. A., and Levin, S. (1997). The MultifacetedAspects of
Knapp, A. K., Blair,J. M, Briggs,J. M., Collins, S. L., Hartnett,
ConservationEcology 1(1): 3. http://www.
C, Johnson,L. C, and Towne,E. G. (1999). The KeystoneRole
EcosystemIntegrity.
of Bison in NorthAmericanTallgrassPrairie.Bioscience 49(1):
1/iss1/art3/.
consecol.org/vol
39-50.
and Ecology: An Approachto
Ellen,R. F. (1978). Nuaulu Settlement
Relationsof an EasternIndonesianCommunity. Kotliar,N. B. (2000). Applicationof the New Keystone-Species
theEnvironmental
The Hague.
Martinus
Conceptto PrairieDogs: How Well Does It Work?Conservation
Nijhoff,
Biology 14(6): 1715-1721.
Ellen, R. F. (1982). Environment,Subsistence and System: The
CambridgeUniversity Lawton, J. H., and Jones, C. G. (1995). Linking species and
Ecologyof Small-ScaleSocial Formations.
Press,Cambridge.
ecosystems:organismsas ecosystemengineers.In Jones,C. G.,
and Lawton, J. H. (eds.), Linking Species and Ecosystems.
Ellen,R. F. (2003). On theEdge of theBanda Zone: Past and Present
in the Social Organisationof a Moluccan Trading Network.
Chapman& Hall, New York,pp. 141-150.
of Hawaii Press,Honolulu.
Levey, D. J. (1990). Habitat-DependentFruitingBehavior of an
University
and Tropical Treefall
of Metroxylonsagu and the
UnderstoryTree, M/comYz-Centrodesma,
Ellen, R. F. (2004a). The distribution
historicaldiffusionof a complex traditionaltechnology.In
Gaps as KeystoneHabitatsforFrugivoresin Costa Rica. Journal
of TropicalEcology 6(4): 409^420.
Boomgaard,P., and Henley,D. (eds.), Smallholdersand Stockbreeders:Historiesof Food Crop and Livestock Farmingin
Meilleur,B. A. (1994).. In searchof 'keystonesocieties'.In Etkins,N.
L. (ed.), Eating on the Wild Side. Universityof Arizona Press,
SoutheastAsia. KITLV, Leiden.
Tucson,pp. 250-279.
Ellen,R. F. (2004b). ProcessingMetroxylon
sagu Rottboell(Arecaceae)
as a TechnologicalComplex:A Case StudyfromSouth Central
Menge,B. A., Berlow,E. L., Blanchette,C. A., Navarette,S. A., and
Yamada, S. B. (1994). Variationin InteractionStrengthin a
Seram,Indonesia.EconomicBotany58(4): 600-625.
of
Palm
and
Local
R.
F.
Rocky IntertidalHabitat. Ecological Monographs64(3): 249Management Sago
Ellen,
Knowledge
(2006).
286.
sagu Rottboell)Diversityin South CentralSeram,
(Metroxylon
Miller, B., Reading, R., Hoogland, J., Clark, T., Ceballos, G.,
Maluku,EasternIndonesia.Journalof Ethnobiology26(2): 258298.
List, R., Forrest,S., Hanebury,L., Manzano, P., Pacheco, J.,
and Uresk, D. (2000). The Role of Prairie Dogs as a
Garibaldi,A., and lurner,N. J. (ZUU4a).culturalReystonespecies:
and Restoration.
forEcologicalConservation
Keystone Species: Response to Stapp. ConservationBiology
Ecology
Implications
and Society9(3): 1. http://www.ecologyandsociety.org/vol9/iss3/14(1): 318-321.
artl.
Mills, L. S., Soule, M. E., and Doak, D. F. (1993). The KeystoneSpecies Conceptin Ecology and Conservation.Bioscience43(4):
Garibaldi,A., and Turner,N. (2004b). The Nature of Cultureand
219-224.
Keystones.Ecology and Society 9(3): r2. http://www.ecology
Nabhan,G. P., and Carr,J. L. (eds.) (1994). Ironwood:an ecological
andsociety.org/vol9/iss3/resp2/.
and cultural keystone of the Sonoran Desert. Conservation
G., Kaiser,M. J.,and Castilla,J.C. (2006).
Gelcich,S., Edwards-Jones,
Occasional PaperNo. 1, ConservationInternational,
International
Policy Can Reduce Resilience in Traditionally
Co-management
WashingtonD.C.
ManagedMarineEcosystems.Ecosystems9: 951-966.
O'Neill, R. V, and Kahn, J. R. (2000). Homo Economus as a
Grimm, N. B. (1995). Why link species and ecosystems/ A
KeystoneSpecies. Bioscience 50(4): 333-337.
perspectivefrom ecosystem ecology. In Jones, C. G., and
Paine, R. T. (1966). Food Web Complexityand Species Diversity.
Lawton,J.H. (eds.), LinkingSpecies and Ecosystems.Chapman
AmericanNaturalist100: 65-75.
& Hall, New York,pp. 5-15.
resourceuse, conservationand
T. B. (2002). Ethnoecology,
Paine, R. T. (1969). A Note on TrophicComplexityand Community
Henfrey,
in theSouthRupununi,
Stability.AmericanNaturalist103: 91-93.
developmentin a Wapishanacommunity
http:// Payton,I. J., Fenner,M., and Lee, W.Ci.(2002). KeystoneSpecies:
Guyana. Ph.D. thesis,Universityof Kent at Canterbury,
The Concept and Its Relevance for Conservation in New
lucy.ukc.ac.uk/csacpub/Henfrey_thesis/.
Zealand. Science forConservation203.
T. B., and Platten,S. (2006). On culturalkeystonecomplexes
Henfrey,
Piraino,S., and Fanelli,G. (1999). KeystoneSpecies: WhatAre We
and theirimplicationsfor systemfunction.In Cyberneticsand
Talking About? ConservationEcology 3(1): r4. http://www.
Systems2006: The Proceedingsof the 18thEuropeanMeeting
1/resp4/.
on Cyberneticsand System research. Austrian Society for
consecol.org/vol3/iss
innovation
Platten,S. J. (2005). The culturaldynamicsof agricultural
Cyberneticstudies,Vienna.
of
Kent.
Thesis.
Ph.D.
in
a
Minahasan
as
Herbivores
Dominant
University
W.
J.
village.
Keystone
(2002). Functionally
Higdon,
Species. ConservationEcology 6(2): r4. http://www.consecol. Platten,S. J. (2007). Carrotsand clove: Traditionalchangein upland
In Ellen,R. F. (ed.), Traditional
Minahasanagriculture.
Ecological
org/vol6/iss2/resp4/.
and
Problem:EcologicalRestoration
Knowledgeand ModernCrisesin IslandSoutheastAsia. Berghahn,
Higgs,E. (2005). The Two-Culture
Oxford.
ofKnowledge.Restoration
theIntegration
Ecology13(1): 159-164.
Power,M. E., and Mills, L. S. (1995). The KeystoneCops Meet in
ueometry,ana
Holling, C. S. (1992). Cross-Scale Jviorpnoiogy,
Hilo. Trendsin Ecology and Evolution10(5): 182-184.
DynamicsofEcosystems.EcologicalMonographs62(4): 447-502.
Hurlbert,S. H. (1997). FunctionalImportancevs. Keystoneness: Power,M. E., Tilman,D., Estes, J. A., Menge, B. A., Bond, W. J.,
Mills, L. S., Daily, G., Castilla,J. C, Lubchenco,J.,and Paine,
ReformulatingSome Questions in Theoretical Biocenology.
R. T. (1996). Challengesin theQuest forKeystones.Bioscience
AustralianJournalof Ecology 22: 369-382.
or
the
from
Structured
Are
E.
46(8): 609-620.
C.
Top-Down
Ecosystems
(1998).
Kay,
Kosemund,A. D., and Anderson,c a. (zuujj. engineeringKoie
Bottom-Up:A New Look at an Old Debate. WildlifeSociety
Models: Do Non-humanSpecies Have the Answers?Ecological
Bulletin26(3): 484-498.
W. G., and Kay, F. R. (1997). Mechanisms
Engineering20: 379-387.
Kerley,G. I. H., Whitford,
Rather
Schulze, E. D., and Mooney, H. A. (1994). Ecosystemfunctionot
forthe KeystoneStatusof Kangaroo Rats: Graminivory
Than Granivory?
biodiversity:a summary.In Shulze, E. D., and Mooney,H. A.
Oecologia 111(3): 422^28.
} Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Hum Ecol (2009) 37:491-500

500
(eds.), Biodiversityand EcosystemFunction.Springer,Berlin,
pp. 497-510.
Simberloff,D. (1998). Flagships, Umbrellas, and Keystones: Is
Single-Species Management Passe in the Landscape Era?
Biological Conservation83(3): 247-257.
St. Antoine,S. (1994). lronwoodand art:lessons in culturalecology.
In Nabhanand Carr(eds.), lronwood:An Ecological and Cultural
Keystone of the Sonoran Desert. ConservationInternational
Occasional PaperNo. 1, ConservationInternational,
Washington
D.C., pp. 69-85.
Stercho,A. M. (2006). The Importanceot Place-BasedFisheriesto the
KarukTribeof California:A SocioeconomicStudy.M.A. thesis,
HumboldtStateUniversity.
Steward,J. H. (1955). Theory of CultureChange: The Methodology of Multilinear Evolution. Universityof Illinois Press,
Chicago.
Stoffle,R. W., Halmo, D. B., Evans, M. J.,and Olmsted,J.E. (1990).
CalculatingtheCulturalSignificanceof AmericanIndianPlants:
Paiute and Shoshone Ethnobotanyat Yucca Mountain,Nevada.
AmericanAnthropologist
92(2): 416-432.
Tanner,J. P., Hughes, E. P., and Connell, J. H. (1994). Species
Coexistence,Keystone Species, and Succession: A Sensitivity
Analysis.Ecology 75(8): 2204-2219.

J.(1986). Keystoneplantresourcesin thetropicalforests.In


Terborgh,
Soule, M. (ed.), ConservationBiology: The Science of Scarcity
and Diversity.Sinauer,Sunderland,pp. 330-344.
The Snow Leopard Conservancy.(2007). MountainCultures,Keystone Species: Exploringthe Role of CulturalKeystoneSpecies
in CentralAsia. Final Report(Grant2005-2019) submittedto
The ChristensenFund by SLC/ Cat Action Treasury,Sonoma,
California.
Ticktin,T., Amaka Whitehead,A. N., and O'Ala Fraiola,H. (2006).
TraditionalGatheringof Native Hula Plants in Alien-Invaded
Hawaiian Forests:AdaptivePractices,Impactson Alien Invasive
ConservaEnvironmental
Species and ConservationImplications.
tion33(3): 185-194.
Turner,N. J. (1988). 'The Importanceof a Rose': Evaluatingthe
CulturalSignificanceof Plantsin Thompsonand LillooetInterior
Salish. AmericanAnthropologist
90(2): 272-290.
CarbonLock-In.EnergyPolicy
Unruh,G. C. (2000). Understanding
28(12): 817-830.
Vanclay,J. (1999). On theNatureof KeystoneSpecies. Conservation
Ecology 3(1): r3. http://www.consecol.org/vol3/issl/resp3/.
Willson, M. F., and Halupka, K. C. (1995). AnadromousFish as
Keystone Species in VertebrateCommunities.Conservation
Biology 9(3): 489-497.

^y Springer

This content downloaded from 130.238.7.43 on Wed, 5 Nov 2014 09:25:24 AM


All use subject to JSTOR Terms and Conditions

Вам также может понравиться