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Contributed

TheUseofSpeciesAccumulation
forthePrediction
Functions
ofSpeciesRichness
JORGESOBERON M.
Centrode Ecologia
UniversidadNacional Autonomade Mexico
Apdo. Postal 70-275
Mexico D. F.

JORGELLORENTEB.
Facultadde Ciencias
UniversidadNacional Autonomade Mexico
Apdo. Postal 70-399
Mexico D. F.

Abstract:We develop a stochastic theoryof the accumula-

tion of new species in faunistic orfloristicinventories.Differentialequationsfor theexpectedlistsize and its variance


as a function of the timespent collectingare presentedand
solvedforparticularcases. Theseparticular cases correspond
models of bow theprobabilityof adding a new
to different
species changes withtime,thesize of thelist,thecomplexity
of thearea sampled, and otherparameters.Examples using
and mammals are discussed,and
field data from butterflies
it is argued thattheequations relatingsampling effortwith
size of the list may be usefulfor conservationpurposes because theyshould lendformalityto comparisonsamong lists
and because theymay have predictivepower by extrapolating theasymptoticsize of thelists.Thesuitabilityofdifferent
models to a varietyoffield situations is also discussed.

Introduction
Faunisticand floristicstudies oftenreveal that as the
timespentcollectingincreases,the numberofnew spePaper submittedSeptember26, 1991; revisedmanuscriptaccepted
March 9, 1992.
480
ConservationBiology
Volume 7, No. 3, September1993

El uso de funciones
de acumulacion
de especiesparala
prediccion
de la riguezade especies
Resumen: Se presenta una teoria estoca'sticade la acumulacion de especiesnuevas en inventariosfloristicos
ofaunisticos.Se obtieneny resuelvencasos particulares de las ecuaciones diferencialesque relacionan el tamafio esperado de
las listasy su variancia con el tiempodedicado a la colecta
Los casos particulares correspondena diferentesmodelos de
como laprobabilidad de aniadiruna especie nueva a la lista
cambia con el tiempo,el tamafiode la lista; la complejidad
del area y otrosparacmetros.
Se discutenejemplos con datos
de campo de mariposasy mamiferosy se argumenta que el
contar con ecuaciones que relacionen el esfuerzode colecta
con el tamafiodel inventariopuedeser uztilparapropositos
conservacionistasporque se podran formalizar las comparaciones entre inventarios y porque tales ecuaciones
pueden tenerun valorpredictivoal extrapolarpara obtener
los valores asint6ticos de las listas. Tambie'nse discute la
conveniencia de los diferentesmodelos a distintas situaciones de campo.

cies added to the list asymptotically


approaches some
ceiling.In a paper on inventoriesof butterfly
species,
Clench (1979) proposed the use of the MichaelisMentenequationto describeempiricallythebehaviorof
thecummulativespecies-effort
relationship.Despite the
potentialutilityof such a relationship,lepidopterists
have only recentlybegun to use it (Lamas et al. 1991;

& Ilorente
Soberon

Prediction
ofSpeciesRichness 481

Raguso & Llorente 1993). NeitherClench's nor other


relatedequations are commonlyused in faunisticstudies. In at least one botanic paper, Miller and Wiegert
(1989) have used a related equation (an exponential
model; see below) to predictthe totalnumberofplant
species expected in a region.Althoughthe use of such
functionsis still uncommon,it is more widespread in
plottingspecies versuseffort
to estimatevisuallywhether an asymptotehas been reached (Miller & White
1986; Milleret al. 1987; Miller& Wiegert1989; Newmark1991).
Havinga theoreticalbasis forunderstanding
the relationshipbetweencollectingtimeand numberofspecies
accumulated would be useful because, among other
things,(1) it would give formality
to faunisticand floristicwork by allowingmore rigorousand quantitative
comparisonsbetween lists,(2) itwould providea planningtool forcollectingexpeditions,and (3) it mayprovide a predictivetool forconservationand biodiversity
studies,ifused to extrapolatethe totalnumberof species presentin an area. In thispaper,we presenta stochasticmodel oftheprocess of addingnew species to a
listand we will derivesolutionsfordifferent
biological
situations.For one of thesewe obtainthe variance,the
lackingofwhich,as pointedout by Lamaset al. (1991),
is a drawbackofClench'smodel.We fita numberofdata
sets to our equations and discuss the usefulnessand
limitationsof thismethod.

TheModel
A simple model of the process of accumulatingnew
species is the pure birthprocess (Bailey 1964; Pielou
1969). This model assumes that the systemis represented by states,in our case the numberof different
species,and thata suitabletimeincrementmaybe chosen such thatthe systemeithermoves to the next state
or remainswhere it was at time t. In symbols:
prob(ij
prob(i j)

+ l)At = X(,t) At
&t = 1 -X(j ,t)At.

(1)

In words,the probabilityof addingone species to a list


of size j in the time intervalAt is denoted by X(j,t)At,
andwe assumethetimeintervalis so smallthattheonly
otherpossibilityis thatin thesame timeintervalno new
species is found,withprobability1 - X(j,t)At.The symbol X(j,t)At denotes the probabilityof adding a new
species to the list,aftera collectingtime At and given
thatwe alreadyhavej species in timet. The expression
HenceX(j,t)Atis a per-unittimetransitionprobability.
forthwe shallreferto X(j,t) as thecollectingfunction.It
should be clear that the particularshape of X(j,t) depends on factorssuch as the samplingmethod,the size
of the area sampled,and coverage of suitablehabitats.

The collectingfunctionis a functionbothofthe biology


of the taxon of interestand of the methodsused.
Withthis definition,
we now ask for the probability
P(J)tthatat timet the listhas exactlyj species (this is a
stateprobability).It is shownin textbooks ofstochastic
processes that such a probabilityobeys the following
equations:
dp(j)t/d t = p(i

1)t X(i - 1,t)

P(i)t X(jt).

(2)

It is importantto make the technicalpoint thatsince


the onlypermittedtransitionis fromj1 to j in the
time unit At,Equations 2 are a particularcase of the
nonhomogeneousand more generalKolmogorov'sforward equations (Bailey 1964: 77), and it is allowed to
have X as a functionof time.
The above set of equations (one foreach -state
j), if
solved,will yield the distribution
ofprobabilitiesof the
size of the list at time t Althoughthe systemcan be
solved forparticularmodels X(j,t), and purpose of this
paper requiresonly expressionsfor the expected size
and varianceof the list.Aftersome algebra(outlined in
theappendix) itcan be shownthatthedifferential
equationsforthe firstand second momentsof the distribution ofj at time t are simply
d (/)/dt = I pi A(j,Xt)
d ()/d t = 2 12jpt(j) X(j,t) +

(3)
ptj) X(,t), (4)

where the sums are takenfromj = 0 to infinity.


Generatlyspeaking,aftersubstitution
ofparticularmodels of
the collectingfunctionX( ,t) in Equations 3 and 4, we
solve the differential
equationsand obtainthe expected
value of the numberof species in time t: E(j ,t) = (j)t
henceforthdenoted as S(t), and its variance V(j,t) =
2)t- (_i)2t. A numberof interesting
quantitiescan, in
principle,be derivedfromthe moments.For example,
the list size for long times,the time to accumulate a
certainfractionof the asymptote,the timeto lower the
per capita rate of species increase below a certain
threshold,and the confidencelimitsfollow fromthe
solutionsto Equations3 and 4. In the followingsection
we shallfindsome of these forparticularcases.

LinearDependenceon j
The simplestcase is when the collectingfunctiondepends linearlyon the size of the listand the parameters
are constantin time:
X(j,t)= a - b j,'

(5)

meaningthatas the species listgrows,theprobabilityof


adding a new species to the list in the intervalAt decreases proportionallyto the currentsize of the list,
eventuallyreachingzero. This model may be adequate

ConservationBiology
Volume 7, No. 3, September1993

482

ofSpeciesRichness
Prediction

Soberon& LUorente

when one is samplinga relativelysmallarea, or a wellknowngroup,or both,and eventuallyall species will be


registered.
The expressionsforthe mean and varianceobtained
by substitutingEquation 5 in Equations 3 and 4 and
solvingthe differential
equations are
S(t) = a/b[ 1 - exp( - b t)]

(6)

V(t) = S(t) exp( - b t).

(7)

The parametera representsthe listincreaserate at the


beginningof the collection,and the asymptoteis given
bya/b.a has unitsofspecies X time- l, and b oftime- l.
Both parameterscan be obtained by nonlinearregression procedures,and below we shallassume thata suitable algorithmis availablewithoutenteringintothe detailsof nonlinearfitting.
Lamas et al. (1991) asked forthe timetq requiredto
registera proportionof the totalfaunaq = SIR, where
R = a/b representsthe asymptoteor total richnessof
the site. FromEquation 6, thisis simply
tq =

-l/b ln(1 - q).

(9)

For example,how long it will take to reach 90% (q =


0.9) of the asymptoticsize ofa listifb = 0.1 per week?
Equation9 gives the answeras 23 weeks. Althoughthis
resultis interesting,
given the asymptoticbehaviorof
Equation 6, reachinga 100% richnessrequiresan infinitetime.It maybe more usefulto ask how long it will
takefortherateofper capitaspecies increment(dS/Sdt)
to go below a particularsize. For example,how much
collectingtimeit is requiredfordS/Sdt < 0.01? Calling
the thresholdk (which has unitsof time-1), the time
needed to lower the per capita listincreaserateis simply:
tk

= l/b ln(l + b/k).

species collected at timetk Substituting


Equation 10 in

Equation6, we obtainS(tk)

a/(b + k), andthecor-

Exponential
Dependenceon j.
A slightlymore complex model forthe collectingfunction ariseswhen we assume thatincreasingthe size of
thecollectiondecreases theprobabilityofaddinga new
species in a nonlinearway. The simplestsuppositionis
an exponentialdecrease:

(11)

X(jt) = a exp (-b j).

This model may be reasonable in cases in which the


regionbeingsampledis largeor the taxa poorlyknown,
and thusthe probabilityof findinga new species never
reaches zero.
Substitutionof Equation 11 in Equation 3 yields an
equation that can be solved by noting that I p(1)
oftheprobability-generating
exp( - bj) is thedefinition
function(PGF) of the distribution
p(j). By postulating
different
we can solve the equation.A readistributions,
sonable assumptionis thatthep(j) are Poisson distributed,withPGF = exp( -z ()) and z = 1 - exp( - b).
Then it is possible to obtainthe expectationS(t), which
is simply:
S(t) = 1/zln(1 + zat).
Anothercomplicationariseswhen we have exponentiallydecreasingprobabilitiesof adding a new species,
but allow themto reach a value of zero:
X(j,t) = a exp(- b j) - c.

(10)

If,forexample,b = 0.1 per week, and k = 1%, thentk


- 24 weeks,meaningthat24 weeks afterthebeginning
the listwill be growingat 1% of the currentsize, per
week.
Since the standarderrorofS(t) is the square root of
the variance,we can use Equation 7 to estimateconfidence limitsof a givenspecies count. In particular,we
can estimateconfldencelimitsforS(tk), the numberof

respondingstandarderroris (a k)112/(b + k). In the


absence of a fullprobabilitydistributionforS(t), it is
possible to use the rule of thumb that two standard
errors approximate a 95% confidence interval.The
above resultsare summarizedin Table 1.

(12)

Again,the expectationS(t) can be obtained:


S(t) = l/z ln [a/c - (a - c) exp( - czt)/c].

The ClenchEquation
The Michaelis-Menten
equation used by Clench (1979)
can be derivedfromthemodelpresentedhere,bygoing

ofthecollecting
functions
discussed.
Table 1. Somestatistics
Exponential
,(Jt)
S(t)
Var(t)
tq
tk

Asymptote

a-bj
ab(1 - ebt)
S(t) ebt
l/bln[1/(1 - q)]
1/bln(1 + b/k)
a/b

ConservationBiology
Volume 7, No. 3, September1993

Logaritbmic
ae bi
lz ln(1 + zat)
k - za/[(1 + zat) ln(1 + zat)]

Clencb
a + b/a[S(t)2 - 2j
at/(1 + bt)

q/[b(l - q)]

[(1 +
a/b

4bk)1/2

1J/2b

& Llorente
Soberon

Prediction
ofSpeciesRichness 483

backwardson the derivationto obtain its implicitcollectingfunction:


S(t)2
X(j,t) = a + b2/la

2 bj/ a

(13)

2bj/a

(14)

or, equivalently,
X(j,t) = a + b2/a[at/(l

+ bt)]2

Substituting
eitherofthe above equationsin Equation3
and solvingyields
S(t)

a t/(l + b t),

which is Clench's equation with a slightlydifferent


parametrization.
In Equation 13, the expectationappears
in the collectingfunction,therebyincreasingits value.
in Equation 14, fora given value ofj the colSimilarly,
lecting functionis larger if the time accumulated is
larger.Biologically,this means that the probabilityof
adding new species will improve(up to a ceiling) as
more timeis spentin the field.This seems to be a very
plausiblemechanism.It makessense to suppose thatas
one accumulates experience with the site, taxa, and
methods,the chances of adding new species will improve.It is veryinteresting
thatClench's equation,originallyproposed only on empiricalgrounds,appears to
have a sensible theoreticalbasis.
Otherparticularcases can be solved:forexample,an
exponentialcollectingfunctionwith negative-binomial
distributionof the p(j)s, and some time-varying
functions.Clearly,each set of assumptionsabout collecting
functionswill yield different
predictionsof the size of

the species accumulationin inventorystudies.We will


proceed to fitsome data sets and to discuss the results.

Examples
Lamas et al. (1991) presentdata obtainedfroma 200person-hourscollection (during September 1989) in
the Pakitza biological station,Parque Nacional Manu,
Madrede Dios, Peru.Theyfittheirdata to the equation
of Clench and obtaineda verygood fit.They also estimated the asymptote(905 species) and calculated the
time required to reach different
percentagesof it. We
digitizedthe information
fromtheir Graph 1, and in
Figure1 and Table 2 we presenttheresultsoffitting
the
Clench,the exponential,and the logarithmicfunctions
to datafromLamaset al. ( 1991). The modelswere fitted
by the quasi-Newtonmethodprovidedby the package
STATISTICA(StatSoft1991).
It is clear thatalthoughthe data fitwell to each ofthe
functions,
theyextrapolateto verydifferent
numbersof
species. In fact,it is impossibleto choose the best ofthe
threemodelsbased solelyon the data set. To choose an
equation,one has to decide whichunderlying
collecting
model describes most accuratelythe particularsituation.For the threeequationsdiscussed above the models are (1) the probabilityof adding new species decreaseslinearlywiththesize ofthelist(Equation 5); (2)
addinga new species becomes more and more difficult,
butneverreacheszero (Equation I 1); and (3) theprobabilityof addinga new species eventuallyvanishes,but
fieldexperience increasesit (Equation 13).

900
800
700
600
a 500
C', 400

300
........
....;.-

20 0

O~~~~~~~~~~~~~
0

100

200

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

300

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

400

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

500

600

700

Person-Hours
Figure 1. Accumulation curvefor butterfliesin Pakitza Peru Data fromLamas et al. (1991). a correspondsto
the logarithmicequation, b to Clench's equation, and c to theexponential equation.

Conservation Biology
Volume 7, No. 3, September 1993

484

& liorente
Sober6n

ofSpeciesRichness
Prediction

ofthefourexamples.
statistics
Table 2. Regression
Atoyac2

Pakitza'
r.2

0.99
7.57
0.0085
890

0.96
6.64
0.0134
495

r.2

0.99
6.72
0.011
611

0.95
5.63
0.0155
363

a
b
asymptote

Clench

a
b
asymptote

Exponential

0.97
8.413
0.00675

0.99
8.869
0.00347

r.2

a
z

Logarithmic

Powdermilt'

Chajut3
0.967
2.88
0.035
82

0.986
1.073
0.0135
79

0.972
2.65
0.047
56

0.988
0.761
0.011
69
0.917
2.447
0.065

0.96
3.207
0.0356

Time units:
' person-hours '.
2person-days'.
3 nights '.
4person-hours '.

In thecase ofthe data ofLamaset al. (1991), it seems


likelythatthelog model (Equation 11) will be adequate
to extrapolate,giventhesize ofthearea,thecomplexity
ofthe fauna,the factthatthe listsize is stillfarfromthe
asymptote,and the yearlyfluctuationsmany tropical
species undergo.All these pointssuggestthat
butterfly
the probabilityof findingnew species will stillbe differentfromzero aftera sizeable increase of the collection effort.It is interestingto extrapolatethe models
fittedto the first200 hours of data to the list size that
Robbins(personal communication)has reportedafter
565 person-hours.At thattime the Clench model predicts737 species and the log model 839, while the true

value is 979. Althoughthisextrapolationcovers an increase of more than 180% over the time intervalused
the models,and thisintervalincludes a nonforfitting
partofthe curve,thelog model predictsthe
asymptotic
correctvalue whithin15%.
In anothercase, Vargaset al. (1991) reportedtheir
sampling,over threeyears,of a large transect
butterfly
(300-2500 metersabove sea level) fromsemideciduous rainforestto pine forest.In Table 2 and Figure2,we
themodels.As in thePakitza
presenttheresultsoffitting
data, the three models provide excellent fittingsin
termsofexplainedvariance,displaysimilarresidualdisbut predictcontrastinglong-termbehavior.
tributions,

600

500

a) 400

2~00

o~~~~~~~~~~~~~~~~

00

100

200

300

400

500

600

Person-Days
Figure2. Accumulation curvefor butterfliesin Sierra de Atoyac,Mexico. From Vargas et al (1991). a corresponds to the logarithmicequation, b to Clench's equation, and c to theexponential equation

ConservationBiology
Volume 7, No. 3, September1993

Prediction
ofSpeciesRichness 485

& Llorente
Sober6n

As in thepreviousexample,and forsimilarreasons,it
is reasonableto assume thateitherthe log or Clench's
model maybe betterpredictorsof the futurebehavior
Theypredicta species increaseof
ofthesamplingeffort.
around 15% (Clench) or 20% (log) ifsample effortis
doubled to 300 person-days.The exponentialmodel,on
the otherhand,predictsan increase of only about 1%
which in thiscase seems unafterdoublingthe effort,
likelysmall.
Anotherexample is the listofbat species reportedby
Medellin(1986 and unpublished)fromthe Chajul Biological Stationin the Lacandon rainforestof southern
Mexico. The collectionsof bats have been ongoingfor
about seven years,usingmistnets at severalspots near
the station.Table 2 and Figure3 show the results.As in
the previous cases, variance explained by each model
and residuals are very similar.Accordingto Medellin
(personal communication),his methodsare well establishedand thearea,althoughveryrich,is relativelysmall
(a few hundred hectares), so the exponentialmodel
should apply.This predicts an increase of about 10%
afterdoublingthe samplingtime.
In our last example we reanalizedthe example provided by Clench (1979) to illustratehis species-time
formula.This studywas carried out for 13 years and
totaled820 hoursofcollectingand observingbutterflies
at the 2000-acre PowdermillNature Reserve in WestmorelandCounty,Pennsylvania.The list appears to be
withonlyone species added in
almostin theasymptote,
the lastfouryearsofdata. The resultsappear in Table 2
and Figure4. The nonlinearfitto Clench's model gives

an asymptoteof79 species. Clench (1979) did notspecifyhis fitting


method,which yieldsan asymptoteof 78.
However,he suggestsa simplified
methodbased on eyefitting
a curveto the data.This is unreliable,as we have
seen that very different
predictionscan be obtained
fromfittinga varietyof models. By doubling the samplingtimeand usingClench's model,an increaseof 4%
of the list is predicted.The exponentialmodel, which
assumesa lineardecrease of the probabilityof addinga
new species, should not be as good a model for the
sampling of butterflyfauna because temperate lepidopteranspecies are knownto undergomarkedabundance cycles (see Taylor& Taylor1977), and therefore
the probabilityof addingnew species should decrease
slowerthanlinearly.The fitofthe exponentialmodel to
Clench'sdata illustratesa problemraisedby Lamaset al.
(1991): the estimatedspecies richnessis smallerthan
the last data point. This is due to the very quick approach to the asymptotethat characterizesthe exponentialmodel.As Lamaset al. ( 1991) state,thisproblem
can be overcomeby fitting
the data with a highweight
assignedto the last point.

Discussion
The use ofextrapolationsofspatialdata to estimatespecies richnessis not new (Kernshaw 1973; Palmer1990,
among others) and can be traced back to the classical
works of Preston(1948, 1962a, 1962b), Fisher et al.
(1948), and others.To our knowledge,however,extra-

80
70
60
50
CI)~~~~~~~D
G)~

40
3

~ ~ ~

0 k
1

u 04

....'......

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

20

40

'.

'.

60

80

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

100

120

Nights

Figure3. Accumulation curvefor bats in Cbajul, Mexico. From Medellin (1986 and unpublished). a corresponds to the logarithmicequation, b to Clench'sequation, and c to theexponential equationr

ConservationBiology
Volume 7, No. 3, September1993

486

& Llorente
Soberon

ofSpeciesRichness
Prediction
100

90
80

70
(

60

a,

50

tX

50

40

I,X

40
20

250

750

500

1000

1250

1500

Person-Hours
Figure 4. Accumulation curvefor butterfliesin PowdermillReserve,Pennsylvania Data from Clench (1979). a
correspondsto the logarithmicequation, b to Clench's equation, and c to the exponential equationo
apolationsover the timedomainhave not been systematically pursued by conservation biologists (Clench
1979; Miller& White 1986; Lamas et al. 1991).
One of thepotentialuses of such methodologycould
be to lend rigorto faunalinventoriesof areas.In poorly
collected sites,which oftenare importantforconservation purposes,reportinga numberof species may be
misleadingwithout some informationabout how far
fromcomplete such lists are. Eitherthe rates of accumulationof new species or an estimateof the percentage ofthe totalnumberis necessaryto makemeaningful
comparisons.Obviously,a place in which 80 species of
have been reportedwith 0.1 additionalspebutterflies
froma place withthe
is verydifferent
cies/person-hour
same 80 species and a rate of 0.01 additionalspecies/
person-hour.In orderto make such comparisonspossi(time/personand numberof persons) alble, the effort
located to the addition of new species should be
reported.It is clear,however,thatas with the size and
ofpersonsofdifferent
compositionof the list,the effort
expertisemay not be equivalent,thus hinderingcomparisonsbetween lists.It is also clear thattimein itself
is notwhatcounts,but how thistimeis distributedover
the seasons. For example 50 person-hoursduringthe
fromthe same 50 perdryseason maybe verydifferent
son-hourswell distributedover one year. We shall returnto thispoint later.
Predictingtherichnessofthefaunaofa site,giventhe
We
known accumulationcurve,would be interesting.
believe thatthe models presentedhere can be used to
thispurpose with some precautions.First,a sample biased eithertemporallyor spatiallyis useless forextrap-

ConservationBiology
Volume 7, No. 3, September1993

olation. For example, collectingonly duringthe rainy


edges,or canopy of a
season,or onlyin the understory,
forest,will yieldextrapolationsvalid onlyforthe spatial
and temporalconditionssampled.The curvesaggregate
variationin the taxa, the samplingmethods,and the
the organspatialand temporalheterogeneityaffecting
isms,and extrapolationsmusttake thisinto account.
Second,choosingan adequate model ofthecollecting
methodsis criticalto accurateestimationoffaunalsize.
in theirextrapomodels divergesignificantly
Different
exceedinglywell to the same set of
lationswhile fitting
data. In thispaper we have used threemodels,but the
generaltheorypresentedallows the derivationof accumulationcurvesfora varietyof collectingfunctions.In
choosinga suitablemodel,theresearcherneeds to state
explicitly its underlyingassumptions. Because this
choice is to some extent subjective,developing more
objectiveproceduresforchoosinga model should be a
priority.
models requiresinforChoosingamong the different
mationabout the size of the area sampled and the kind
of faunaor florain question.One extremecase is samplingwell-knowntaxa in small or homogeneous areas
with few rare species. In this case, the exponential
model maybe suitable.The otherextremeis sampling
unknown taxa in large or heterogeneous areas with
manyrare species. The Clench or logarithmicmodels
maybe adequate forthese situations.
Clearly,there must be a relationshipbetween the
sampledarea,thespecies-abundancecurve,and thecollectingfunction.Severalauthorshave advanced in this
direction.For example,Millerand Wiegert(1989) gen-

& Liorente
Soberon

erated species-abundancerelationswith canonical lognormal,uniform,


random,and observed extantspecies.
Then they obtained the accumulation exponential
fromthesedata sets.Both
curvesby computer-sampling
the asymptote(a/b) and the increase rate of list size
nearthe origin(a) appear to be similarforthe different
but there are differspecies-abundancedistributions,
ences in the middle part of the accumulationfunction
distributions
sampledfromdifferent
(Miller & Wiegert
1989). Froma different
pointofview,Efronand Thisted
(1976) developed a methodfor the estimationof the
numberof new species thatwill appear aftersamplinga
timeunit.Unfortunately,
theirmethodrequiresthespecies-abundancedistribution
as obtainedby samplingthe
"fauna"duringa previoustime unit,which is difficult.
Finally,an anonymousrefereepoints out that a logseries distribution(Pielou 1969) of species-abundance
in which the numberof individualssampled increases
linearlywithtimewill yield the logarithmicmodel (Table 1). This interestingsubject presentssome difficulties and will be addressedin a futurepaper.
Anotherapplication of the accumulationfunctions
may be the planningof fieldcampaigns.By estimating
thenumberofhoursrequiredto add a givennumberor
itshould
percentageofspecies,givena previoushistory,
be feasibleto estimatecosts of fieldwork in a rigorous
way.Not onlymightthismakepossibletheestimationof
the cost of addingnew species to the list,but because
neartheasymptoterarespecies are likelyto be theones
being added, it may be possible to obtain some value/
cost estimatefordifferent
periodsduringthecollection.
All the curves fittedpresenta veryregulardistribution of residuals.This indicates systematicdeparture
from the assumptionsof regressions.Also, the data
points are not independent.These two points,strictly
speaking,invalidatestatisticalinference,but this is a
pointof statisticalfinessethatmaybe irrelevantforthe
purposes of thispaper. Normally,the biologisttendsto
assess the totalrichnessof a site by extrapolating
from
his or her experience of the place, methods,and taxa,
withoutassigninganyprobabilityof errorto the figure.
The methodpresentedhere is a way to add objectivity
and rigor to such informalpractices. If only because
themethodspretheyexpose theirhiddenassumptions,
sentedare interesting.
More experiencewiththe methodologyand further
developmentofthetheory-in particularits statisticalaspects-will be requiredto decide
whethertheyare usefulforpredictionor planning.

Acknowledgments
We thankGabrielaJimenezforher help withthe graphs
and tables.Dessie Underwoodgave us some helpfuladvice. Town Peters made severalveryusefulcomments
and improvedour Englishsignificantly.
RodrigoMedel-

ofSpeciesRichness 487
Prediction

lin, Robert Robbins,ArmandoLuis, and Isabel Vargas


allowed the use of some unpublisheddata.
Literature
Cited
Bailey,N. T.J. 1964. The elements of stochasticprocesses.
WileyPublicationsin Statistics,
New York,New York.
Clench, H. 1979. How to make regional lists of butterflies:
Some thoughts. Journal of the Lepidopterists' Society
33(4):216-231.
Efron,B., and R. Thisted.1976. Estimationof the numberof
unseen species. Biometrika63:435-447.
Fisher,R.A.,A. S. Corbet,and C. B. Williams.1948. The relation between the numberof species and the numberof individualsin a randomsampleofan animalpopulation.Journalof
AnimalEcology 12:42-58.
Kernshaw,K A. 1973. Quantitativeand dynamicplantecology
Arnold,London,England.
Lamas,G.,R. K Robbins,and D. J.Harvey.1991. A preliminary
surveyof the butterfly
faunaof Pakitza,Parque Nacional del
Manu,Peru,with an estimateof its species richness.Publicaciones del Museo de HistoriaNatural,Universidadde San Marcos, Peru 40:1-19.
Medellin,R. 1986. Murcielagosde Chajul. B.Sc. Thesis. Facultad de Ciencias,UniversidadNacional Autonomade Mexico,
Mexico.
Miller,R. I., and P. S. White.1986. Considerationsforpreserve
designbased on thedistributions
ofrareplantsin GreatSmoky
MountainsNational Park.Journalof EnvironmentalManagement10:119-124.
Miller,R. I., and R. G. Wiegert.1989. Documentingcompleteness,species-arearelations,and the species-abundancedistributionof a regionalflora.Ecology 70(1):16-22.
Miller,R. I., S. Bratton,and P. S. White.1987. A regionalstrategyforreservedesignand placementbased on an analysisof
rare and endangeredspecies distribution
patterns.Biological
Conservation39:255-268.
Newmark,W. D. 1991. Tropical forestfragmentation
and the
local extinctionof understorybirdsin the easternUsamabara
Mountains,Tanzania.ConservationBiology5(1):67-78.
Palmer,M. W. 1990. The estimationof species richness by
extrapolation.Ecology 71(3):1195-1198.
Pielou, E. C. 1969. An introductionto mathematicalecology.
New York,New York.
Wiley-Interscience,
Preston,F. W. 1948. The commonness,and rarity,of species.
Ecology 29:254-283.
Preston,F. W. 1962a The canonical distribution
of commonness and rarity,
vol. I. Ecology43:185-215.
Preston,F. W. 1962b. The CanonicalDistributionofCommonness and Rarity,vol. II. Ecology 43:410-432.

ConservationBiology
Volume 7, No. 3, September1993

488

Prdicton
ofSpecies
Richness

Raguso,R.,andJ.Llorente.1993. The butterflies


oftheTuxtias
Mts.Veracruz,Mexico, revisited:Species-richnessand habitat
disturbance.Journalof Researchon the Lepidoptera.In press.

Sober6n
& Liorente

Appendix
To deriveEquations3 and 4, we beginwiththe definitions
of thefirst
two moments:

(7) =

StatSoft.1991. CSS: Statisticahandbook,vol. II. StatSoftInc.,


Tulsa,Oklahoma.

(2)

Ejp(p)t
= Xj2p(j)

which have derivatives:

Taylor,L. R., and R. A.J.Taylor. 1977. Aggregation,


migration
and populationmechanics.Nature265:415-420.
VargasF. I.,J.Llorente,and A. Luis. 1991. Lepidopterofauna
de
Guerrero.I. Distribuciony Fenologiade los Papilionoideade
la Sierra de Atoyac. Publicaciones Especiales del Museo de
Zoologia #2. Facultadde Ciencias,UniversidadNacional Aut6nomade Mexico, Mexico.

ConservationBiology
Volume 7, No. 3, September1993

dQf/dt=
d(2)/dt =

j dp()/dt

(Al)

j2 dp()/dt

(A2)

Substitution
ofthevalues ofdp()Idt givenby Equation2 yieldsequationsthatcan be simplified,
in the case ofAl, by addingand subtracting Xp(j -1 )X(f - 1) and thensimplifying
and,in the case ofA2, by
addingand subtracting
termsto completetheexpressionsTp(j-1 Xi
1)2 and :p(j - 1)( - 1)3. Furthersimplification
yieldsEquation
4.

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