Nordic Society Oikos

Evenness Indices: Are Any Admissible?

Author(s): Richard D. Routledge
Source: Oikos, Vol. 40, Fasc. 1 (Jan., 1983), pp. 149-151
Published by: Wiley on behalf of Nordic Society Oikos
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Evennessindices: are any admissible?

Richard D. Routledge,Dept. of Mathematics,Simon Fraser Univ.,Burnaby,B.C., Canada V5A 1S6

Summary.Manyindiceshave been proposedformeasuringthe where pi = the proportionalabundance of species 1.

evenness of a biological community,that is, the degree to
Because of the logarithmicnatureof thisindex,she di-
which abundances are divided equitably among the species
vided it not by s, but by log s. Because s can also be
present.In addition,numerousdesirablepropertiesof even-
viewedas a diversity
ness indices have been proposed. It is proved here that no index,thismeasureof evennessis a
ratio of two diversityindices.This definitionhas been
singleindexcan satisfyeven the mostbasic oftheseproperties.
Potentialsolutionsto thisdilemmaare discussed. extendedto includemore generalratiosof diversity in-
dices (Hill 1973), and minor modifications(Hurlbert
1971, Peet 1974). Somewhatdifferent indiceshave also
been proposed (see, e.g., Lloyd and Ghelardi 1964,
Pielou 1977). Many authors(e.g., Sheldon 1969, Pielou
1977, Engen 1979, Taillie 1979, Alatalo 1981) have
Introduction
proposed desirablepropertiesof evennessindices.En-
The conceptof evennesscan be associatedwithgroups gen (1979) and Alatalo (1981) concludethatno single
of similarspecies occupyingthe same trophiclevel in a index can possess all the desirableattributesproposed
community. Evennessdescribesthedegreeto whichthe fora measureof such a loose conceptas evenness.This
abundances are divided equitably among the species conclusionbegs the question, "Can any index meet a
present.For example,a mixedforestcontainingseveral minimalsetofbasic requirements fora sensiblybehaved,
species, all of which are equally abundant, exhibits usable measureof evenness?" Followingis a statement
maximalevenness.A secondforest,containingthesame of such basic requirements,and an appraisal,withre-
species complement,but dominatedby a singlespecies, spectto theserequirements, of potentialcandidatesfor
is judged to be less even. indicesof evenness.
Pielou (1966) defineda popularindexof evennessas
follows: She noted that among all communitiescon-
taininga fixednumber(s) of species, the diversity,as
Basic requirements
well as the evenness,is highestwhen the species are
equally abundant.This observationled her to define An evenness index (E) is to be definedfor any com-
evennessas the ratioof a diversityindexto an indexof munitycontainingtwo or more species. It mustbehave
species richness. She measured diversity by the sensiblywithrespectto certainadjustmentsof species
Shannon-Wienerindex, abundances.For example,if the total abundance of all
the species in a communityis changed,but the propor-
H' =-X~i=1 pi logpi tionsof thistotalassignedto each species are keptcon-

OIKOS 40:1 (1983) 149

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stant,thenthe evennessshouldremainfixed.That is, E nessofthecollection oforganisms actually
sampled,but
should depend only on the proportionalabundancesof not to draw any inference about some largercom-
the species,(Pl, P2, .*,ps ), not on the combinedabun- munity. It mayalso be acceptableifone is studying the
dance of all the species. evennessof a knowngroupof species,someof whom
Furtherbasic requirementsof E are as follows: maybe eithertemporarily absent,or presentin unde-
tectablequantities.As long as temporarily absent
(Ri) The evenness of a communityshould be de- speciesareregarded as beingpresent, butinundetecta-
creased by marginallyreducingthe abundanceof ble quantities, thediscontinuity at zerowillnotpose a
the rarestspecies. seriousproblem.(Taillie's,1979,proposeduse ofLo-
renzcurvesforcomparing communities withrespectto
(R2) The evennessshouldalso be decreasedby the ad- theirequitability is also restricted to suchsituations.
ditionof a further, exceedinglyrarespeciesto the The Lorenzcurvecan be drawnonlyifs is known.See
community. Taillie,1979,fordetails.)
Indicesthatdo notsatisfy (R1) or (R2) quantify a
(R3) Evenness should depend continuouslyon the conceptsomewhatdistinctfromequitability. Engen
proportionalabundance of any species. (1979) considerssuchindicesas "dominanceequita-
bilitymeasures".Wouldit notbe morepropitious to
The firsttwo propertiesare naturalinterpretations of startafreshwithan entirely separateconceptlikedomi-
theconceptof evenness,or as Engen (1979) and Taillie nance,or perhapsto go backto Simpson's(1949) no-
(1979) call it,equitability.Consider(RI). Any removal tionofconcentration?
of abundance fromthe rarestspecies is analogous to Considerthefollowing example:We are to compare
removalof wealthfromthe poorest individual.the re- twohypothetical communities. Eachcontains aninfinite
sultingdistribution of abundances,or of wealth,is less number ofspecieswithabundances distributed
accord-
equitable or even. Similarlyfor (R2), if a penniless ingto thegeometric distribution:
migrantwere to move into a region,the distribution of
wealthin thatregionwould become less equitable. In- pi = (1-i)O-1, fori= 1,2,3,....
deed, these two requirementsare containedin Taillie's
(1979) general requirementsthat an index of equita- Forcommunity one,e = 0.50; forcommunity two,f =
bilitypreservethe naturalorderinginducedby the Lo- 0.75. Thusin community one,50% ofthetotalabun-
renz curvesused by economiststo compare wealthdis- dance is pre-empted by thefirstspecies,50% of the
tributionsaccordingto equitability.See Taillie (1979) remaining abundanceis pre-empted by the second
fordetails. species,etc. Whereasin community two, this pre-
The thirdrequirementis essential in most applica- emptionfactoris only25%. Engen (1979), Alatalo
tions involvingsampling from a large, uncensused (1991), and othersarguethatthe"dominance equita-
community.An arbitrarily small errorin the estimated measure"ofthefirst
bility community shouldbe higher.
abundance of a givenspecies should not induce a sub- By covt. t, Taillie's(1979) analogyto thetraditional
stantialerrorin theestimatedevenness.Sheldon (1969) economicinterpretation of equitabilitysupportsthe
noticed that indices which depended directlyon the positionthatbothcommunities exhibitminimal equita-
species richnesssufferedthisdeficiency. bilityin thattheybothcontaininfinitely many,arbit-
No index of eveness can satisfyeven these basic re- rarilyrare (impoverished) species.Nonetheless, the
quirements.(RI) states thatas the proportionalabun- communities differ
substantiallyinthedegreeto which
small
dance of therarestspecies is reducedto arbitrarily the abundancesare preemptedby, or concentrated
yetpositivelevels,the evennessis decreased. Yet (R2) within, themorecommonspecies.Pielou(1977: 309)
impliesthatifthespecies is sufficiently
rare,thentotally has suggested investigatingtheratioof abundancesof
eliminatingits presence (that is, reducingits propor- the commonest to the second,third,or fourthcom-
tionalabundanceto zero), would increasethe evenness. monestspecies. These ratios could be viewed as
It followsthattheevennessmustjump discontinously as measuring therateat whichresources are pre-empted
the proportionalabundance approacheszero (rigorous bythecommonest species,or as measuring therelated
proofin appendix). In the sense thatno index of even- concepts ofdominance orconcentration.Suchconcepts
ness can satisfyall threerequirements,none is admissi- are worthinvestigating as replacements forthe un-
ble. wieldynotionofevenness.

Discussion
At least one of these basic requirementsmustbe sac-
Appendix
rificed.The indices most commonlyused to date sac- Followingis a morerigorous
accountofthemainresults
rificethe continuity
requirement,(R3). Such a strategy in thetext.Let E represent
an indexof evenness.Let
is acceptable when one wishes only to studythe even- thespeciesbe arrangedin decreasingorderof abun-

150 OIKOS 40:1 (1983)

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dance, and let ai = the absoluteabundanceof species i.
It is convenientto definea- for all i = 12,3... The
species richness(s) is then the largestvalue of i for
whichai > 0. Let A = ESl ai = the totalabundanceof
all species together,and Pi = a1/A= the proportional
abundanceof species i. Furtherlet A, = A - as = the
totalabundanceof all species excludingthe rarest.E is
to be a functionof (PAIP2,...). Let the abundanceof the
rarestspecies be alteredfroma, to b. Then its propor-
tionalabundanceis alteredfromps to q = b/(As+b).
Furthermore, the proportionalabundance of any other
species (i) is altered from pi = ai/(A?+ as) to ai/
(A,+b). This latter expression can be reduced by
straightforward algebrato pi(1-q)/(l-p.). Then let f(q)
representthe evenness of this adjusted set of propor-
tional abundances.We requirethe following:
(RI) f(q) is to be an increasingfunctionof q forall q
strictly between0 and p.
(R2) f(q) < f(O) forsuitablysmallq > 0. (The version
of (R2) in the textstatedthatthe additionof a
veryrarespeciesshoulddecreasetheevenness.It
is equivalentto demand that its subsequentre-
moval should increasethe evenness.)
even at q
(R3) f(q) shouldbe a continuousfunction,
0.
Theorem:No functioncan satisfyall threeof these re-
quirements.
Food-niche relationshipsamong sympatricpredators:effects
of level ofprey identification
Harry W. Greeneand Fabian M. Jaksic Museum of VertebrateZoology and Dept. of Zoology, Univ. of
California,Berkeley,CA 94720, USA
Summary.Niche overlap and niche breadthwere calculated
twicefor16 setsof vertebratepredators,usinghigh(genericor
specific)and ordinal level prey identification
(LPI). Ordinal
LPI consistentlyunderestimates high LPI niche breadthand
overestimateshigh LPI niche overlap. Ordinal LPI can also
seriouslymisrepresent the rankorderof diet overlapsin com-
munitymatricesand nearestneighborestimatesof guildsize.
Previous conclusionsregardingcommunityphenomena and
optimalforagingin otherorganisms,based on ordinalLPI, are
probablypremature.
OIKOS 40:1 (1983)
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Proof. Select any suitablysmall qO in (Op7P),forwhich
by (R2), f(qO)< f (0). Then by (R1), foranyq < q0,f(q)
< f(qO).Hence, thelimitof f(q) as q->O, ifit exists,must
notexceed f(q0). Therefore,thislimitcannotequal f(O),
and f(q) cannotbe continuousat q = 0.
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Niche breadthand nicheoverlapare widelyused meas-
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papers have addressed theirvalue in estimatingcom-
petition(Colwell and Futuyma 1971, Lawlor 1980) as
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preciatedeffectson these statisticsand on the assess-
ment of higherorder communityphenomena in ver-
151