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Simplifying the Jargon of Community Ecology: A Conceptual Approach

Article  in  The American Naturalist · February 1996


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The University of Chicago

Simplifying the Jargon of Community Ecology: A Conceptual Approach


Author(s): J. E. Fauth, J. Bernardo, M. Camara, W. J. Resetarits, Jr., J. Van Buskirk and S. A.
McCollum
Source: The American Naturalist, Vol. 147, No. 2 (Feb., 1996), pp. 282-286
Published by: The University of Chicago Press for The American Society of Naturalists
Stable URL: http://www.jstor.org/stable/2463205
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Vol. 147, No. 2 The AmericanNaturalist February 1996

NOTES AND COMMENTS

SIMPLIFYING THE JARGONOF COMMUNITY ECOLOGY: A


CONCEPTUAL APPROACH

The fieldsof ecologyand evolutionare frequently criticizedforteleological


arguments and inexactlanguage(Popper1974;Peters1976;Thorpe1986;Mills
et al. 1993;Frazier1994).Whilesomeofthiscriticism maybe misdirected (Steb-
bins1977),imprecise languagehasledtothevirtualsynonymy ofimportant terms
thatshouldretaindistinct usages(see, e.g., Gould1977;Janzen1980;Millset al.
1993).Clarifying ecologicalterminology is importantnotonlybecauseitfacilitates
clear communication but also because it makesexplicitthe assumptionsthat
underlieconceptsanddirectsattention to thosein needofempirical verification.
Herewe focuson fourkeytermsthatarewidelysynonymized, ignored, orother-
wise misusedin theliterature of community ecology:"community," "guild,"
"assemblage,"and "ensemble."Usinga Venndiagram,we provideoperation-
allydefinedand distinct meanings fortheseoftenmisapplied terms.
We approachedthisproblemby considering theways thatresearchers com-
monlylimitthe organismsthattheystudy,and thenwe definedthe resulting
unitsas subsetsofthebiologicalworld.Some researchers restrictthemselves to
studying groupsof phylogenetically relatedspecies; theseare definedby their
respectivetaxa, as arrayedin the classicalLinneannestedhierarchy, and are
represented by set A in figure1. Otherresearchers confinetheirstudiesto a
particular physicalarea; theseare definedby geography alone and are depicted
as setB in figure1. Othersstudygroupsofspecies,without regardto phylogeny
or geographicdistribution, that exploitthe same resource.Such resource-
boundedbutnonphylogenetically circumscribed groupsare represented by setC
(fig.1).
Withthesethreeoverlapping sets-delineatedby phylogeny, geography, and
resources-described,it is a simpleexerciseto match them and theirintersec-
tionswiththeappropriate terms(fig.1). Set A containstaxonomic unitsclustered
accordingto commondescent,frompopulations through moreinclusivegroups
suchas families, orders,and so forth.Mostresearchers recognizethespeciesas
thefundamental unitof taxonomy;a varietyof operationaldefinitions forthis

Am. Nat. 1996. Vol. 147, pp. 282-286.


? 1996 by The Universityof Chicago. 0003-0147/96/4702-0006$02.00.
All rightsreserved.

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NOTES AND COMMENTS 283

GEOGRAPHY RESOURCES

/ /L~~ocal
A
SET B
/Guild\sETC

\ / \ ~Ensembles/

ssembla

\ SET A /

PHYLOGENY

FIG. 1.-Populationsunderstudymaybe dividedintothreedistinct sets:thosedefined


byphylogeny (SET A), geography(SET B), andresources(SET C). In thismodel,ecological
"communities" are merelygroupsoforganisms livingin thesameplace at thesametime.
If we applythismeaning ofcommunity, theintersectionsofthesesetsprovideoperational
definitionsfortheterms"assemblage,""ensemble,"and "guild,"as explainedin thetext.
The intersectionof guildsand taxa denotesan entityforwhichno termexists,butsuch
phylogeneticallyrestrictedgroupsusinga commonresourcein different communities are
generally referredto by a compounddescriptor defining resourceand taxon,e.g., pond-
breeding salamanders.

termhave been advancedin recentyears,suchas thebiological,phylogenetic,


and evolutionary species concepts(reviewedin Frostand Hillis 1990). Set B
contains"communities," whichare simplydefinedas a collectionof species
occurring in thesameplace at thesametime.This is one common(butnotthe
only)definition of a "community" (see appendix;see also Schoener1986).We
chose thisdefinition foritssimplicity so longas a researcher
and flexibility; can
placeboundaries aroundherorhisstudysite,a community can be circumscribed
-easily.The boundariesmay be natural(e.g., serpentine soil communities) or
arbitrary (all organisms withina 1-iM2 plotof lawn).The mainpointis that,in
ordertoconstitute a community, theorganisms understudymustnotbe restricted
furtherbyphylogeny orresourceuse. Otherwise, a moreprecisetermis available
(see below). Set C is based on resourceuse and constitutes a "guild." This is
faithfultoRoot's(1967,p. 335)definition ofa guildas a groupofspecies"without
regardfortaxonomicposition"that"exploitthe same class of environmental
resourcesin a similarway." We interpret "in a similarway" to meanin a con-
sumptive fashion;thatis, use of theresourceby one speciespotentially makes
it unavailableforuse by others.Such resource-bounded butnontaxonomically
circumscribed sets(e.g., foliagegleanersandcavitynesters)havegenerally been
studiedby thecomparative method.

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284 THE AMERICANNATURALIST

Phylogenetically relatedgroupswithina community-the intersection ofsetA


and set B-are aptlytermed"assemblages."Thisconforms to commonecologi-
cal usageandremovesthistermfromsynonymy withcommunity (fig.1). Species
thatsharea commonresourceand occurin thesamecommunity-the intersec-
tionof set B and set C-comprise a "local guild."ThisparallelsRoot's (1967,
p. 335)descriptionofthebirdson hisstudyareaas "local members ofthefoliage-
gleaningguild." Phylogeneticallyrelatedspeciesthatexploitthe same class of
resourcesin a similarfashion,butnotnecessarily in thesame community-the
intersectionof set A and set C- are commonly referredto witha compound
descriptor,forexample,pond-breeding salamanders. Finally,theintersection of
all threesets is an "ensemble"(fig.1) as describedby Istock (1973,p. 535)
as "local, taxonomicallycircumscribedspecies assemblages . . . viewed in an
operationalsenseas collections
ofecologically similarspecies." Although Istock
did notuse thetermguildin describing his notionof an ensemble,we suggest
thatsimilarityofresourceuse was implicitin hischaracterization ofecologically
similarspecies.An ensembleis thusa phylogenetically boundedgroupofspecies
thatuse a similarsetofresourceswithina community. Thisdefinitionconforms
withpriorusage(Istock1973)andeliminates synonymy withassemblage,commu-
nity,and guild.Ensemblesrequiredescriptors clarifyingresource,taxon,and
geography, suchas theensembleofseed-eating Mojavedesertinsects.Giventhe
above definitions,themodelassumesonlythattaxa,geography, and resources
are describedat a commontime.
The advantagesofthisframework are twofold.First,no newtermsare intro-
duced.Second,itprovidesclear,operational definitionsofimportant termsthat
have beenmisusedso egregiously thatpopulartextbooks eitherdo notagreeon
a commondefinition (appendix)or do notprovideone (e.g., Colinvaux1993).In
theprimary literature
one needonlyexaminerecentpapersin anymajorjournal
touncovernumerous incongruences.UsingourVenndiagram, itshouldbe simple
to identify
andcommunicate exactlywhatone is studying. We suggestthatcloser
scrutinyof the ecologicallexiconwillfostermoreprecisecommunication and
expose the assumptions hiddenin colloquialmeanings,therebyfurthering the
advanceofecology.

ACKNOWLEDGMENTS

The ideas presentedhereinarose fromdiscussionsof the EcoLunch group


organizedbyH. M. Wilburat Duke University in 1987.We thankH. M. Wilbur,
J. Leimberger, P. Pearman,K. Sigmon,and all of the otherparticipantsfor
contributing
to ourconfusionaboutecologicalterminology.We thankJ.Antono-
vics,R. Brandon,C. J.Gill,R. G. Jaeger,andH. F. Nijhoutfortheirthoughtful
comments on themanuscript.

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NOTES AND COMMENTS 285

APPENDIX
TABLE Al
DEFINITIONS OF "COMMUNITY" IN SEVERAL ECOLOGY TEXTBOOKS

Set Boundaries Definition Source

Space, time The speciesthatoccurtogether in space and Begonet al. 1990


time.
Space, time,interactions An associationofinteracting populations,usu- Ricklefs1990
allydefined bythenatureoftheirinteraction
or theplacein whichtheylive.
A groupoforganisms thatlivealongsideone an- Tudge1991
other,andin whichthedifferent speciesand
individuals withone another.
interact
Space, time,interactions,
phylogeny A groupofinteracting plantsandanimalsinhab- Smith1992
itinga givenarea.
An assemblageofinteracting plantsandanimals Freedman1989
on a sharedsite.
Group of populationsof plants and animals in a Krebs 1985
- givenplace;ecologicalunitusedin a broad
senseto includegroupsofvarioussizes and
degreesofintegration.
aredirectquotations
NOTE.-All definitions fromtheglossary.

LITERATURE CITED

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ties. 2d ed. Blackwell Scientific,Cambridge,Mass.
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286 THE AMERICAN NATURALIST*

Stebbins, G. L. 1977. The defense of evolution: tautologyor theory? American Naturalist 111:
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J. E. FAUTH*
J. BERNARDOt
M. CAMARAt
W. J. RESETARITS, JR.?
J. VAN BUSKIRK||
S. A. MCCOLLUM#
DEPARTMENT OF ZOOLOGY
DUKE UNIVERSITY
DURHAM, NORTH CAROLINA 27706
SubmittedAugust 12, 1994; Revised May 9, 1995; Accepted June26, 1995

* Equal co-authors;address correspondenceto J. E. Fauth; presentaddress: Departmentof Biol-


ogy, College of Charleston,Charleston,South Carolina 29424-0001.
t Presentaddress: Departmentof Zoology, Universityof Texas, Austin,Texas 78712.
t Presentaddress: Departmentof Environmental,Populationand OrganismicBiology, University
of Colorado, Boulder, Colorado 80309.
? Present address: Center for Aquatic Ecology, Illinois Natural HistorySurvey, 607 E. Peabody
Drive, Champaign,Illinois 61820.
11Present address: Departmentof Biological Sciences, Texas Tech University,Lubbock, Texas
79409-3131.
# Presentaddress: Departmentof Biology, Universityof Michigan,Ann Arbor,Michigan48109.

Associate Editor: Deborah E. Goldberg

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