Preliminary estimates of Lepidoptera diversity from specific sites in the Neotropics using complementarity and species richness estimators

7 Pages·2000·3.5 MB·English

by  Michael G Pogue

JournaloftheLepidopterists'Society
53(2),1999,65-71
PRELIMINARY ESTIMATES OF LEPIDOPTERA DIVERSITY FROM SPECIFIC SITES IN THE
NEOTROPICS USING COMPLEMENTARITYAND SPECIES RICHNESS ESTIMATORS
Michael G. Pogue
SystematicEntomologyLaboratory,P. S. I.,Agricultural Research Service, U.S. DepartmentofAgriculture,
%DepartmentofEntomology, National MuseumofNaturalHistory,
SmithsonianInstitution,Washington, D.C. 20560-0168, USA
ABSTRACT. Lepidopterawerecollectedandspeciesrichnessandcomplementarityoruniquenesswerecomparedbetweentworainfor-
estsites:Pakitza,PeruandBeni,Bolivia.Thetotalnumberofspeciescollectedfrombothsiteswas 1,879ofwhich60weresharedresultingin
acomplementarityof96.8%. Non-parametricequationsandspeciesaccumulationcurvesofHemiceras Guenee (Lepidoptera: Notodontidae)
wereusedtocomparespecies richnessbetweenthree rainforestsites, PakitzaandTambopata, PeruandReservaEthnicaWaorani, Onkone
Gare,Ecuador.Clusteranalysis,usingcomplementarityvaluesforselectedsiteswasusedtodeterminealtitudinalrelationshipsbetweensitesin
CostaRica;relationshipsbetweenforesttypesinBrazil;andfaunaldifferencesamongsitesinwesternAmazoniausingHemiceras.
Additionalkeywords: Biodiversity,Notodontidae,Hemiceras, Chao,jackknife.
Biodiversity as defined by E. O. Wilson (Reaka- taxa around the globe. After a site has been sampled,
Kudlaetal. 1997) is "everything". Biodiversityencom- species richness estimates are used to predict how
passes the genes within a single local population or many species were missed during the sampling pro-
species, the species within a local community, and cess, thus arriving at an estimated number ofspecies
communities comprisingthe diverse ecosytems ofthe basedonthe actualnumberobservedplusthenumber
world. Life on earth is supported by the interactions missed. By using species lists, either generated by
andproductsproducedbyallotherlifeonearth.With- samplingatasite orfrom museum collections, species
outbiodiversitythere wouldbe no life on earth as we composition among sites can be compared. These
knowit. Therefore, it is essential that biologists begin comparisons can then be used in setting policy and
to document and record biodiversity, whether it be makinginformedconservation and managementdeci-
how many species ofinsects are in your backyard to sions.
howmanyspecies oftreesin aforesttohowmanyfor- The goals ofthis paperare 1) to emphasize the im-
esttypes intheworld. portance ofadequate sampling, 2) to assess whether
Anyonewithaninterestinnaturalhistorycanbegin inadequate sampling can still be useful in predicting
to study their local biodiversity and to document it. species richness and complementaritybetween study
One basic element of biodiversity is to know how sites, and3) toprovide amethodofusingcomplemen-
many species are present at aparticular site. This pa- tarity to compare faunal relationships between sites.
perwilloutlinehowanyonecanbegintodocumentlo- To accomplish these goals, I compared overall species
cal biodiversityby gathering data on species richness richness and complementarity of Lepidoptera from
orthe numberofspecies presentat asite ataparticu- two rainforest sites, one in Peru and the other in Bo-
larpointin time. Knowingwhat species are present at livia. In addition, I estimated species richness from
a site is essential because it is the first step in under- species accumulation curves and non-parametric esti-
standing the interactions between the species docu- mators to compare the Hemiceras Guenee (Notodon-
mentedandtheirinteractionswithinthelocalcommu- tidae) faunabetween 2 sites in Peruand 1 in Ecuador.
nityandecosystem. Finally, museum specimens ofHemiceras were used
Presently, no one has much ofan idea exactlyhow for faunal comparisons among sites in Costa Rica,
manyspecies arepresenttodayon earth. Estimates of Brazil, andwesternAmazonia, usingcomplementarity
the numberofspecies worldwide range from 3 to 100 andclusteranalysis.
million (Erwin 1982, 1983, Stork 1988, Hodkinson &
Materials and Methods
Casson 1991, May1992, RavenandWilson 1992). The
studyofspecies richness andcomplementarity, orhow Lepidoptera complementarity in SW Amazo-
different species composition is between sites, is es- nia. Lepidopterawerecollectedattworainforestsites:
sentialto assessingglobalbiodiversitypatterns. Beni, Boliviaand Pakitza, Peru. The Beni studysite at
W
To address the question of world insect diversity 14°49'S, 66°28' is40km E ofSanBorja, at250me-
onemustgetaccurate estimatesofsite-specificspecies ters elevation. Pakitza is located on the Rio Manu at
richness for a variety oftaxa (Colwell & Coddington 11°56'47'1S, 71°17WW within the large drainage
1994), andthen to compare these species liststo mea- basin ofthe Rio Madre de Dios in southeastern Peru,
sure relative levels of overlap and richness of these at356m elevation, approximately550km NWofBeni.
( I I I
66 Journalofthe Lepidopterists' Society
40T
— — — —
t- oHn 1 in1 1 1 I— 1 1 co1 1 m1 1 1 1 1 1 h
cm oj eg
TrapNights
Fig. 1. SpeciesaccumulationcurvesofHemicerasatPakitzaandTambopata,Peru,andOnkoneGare,Ecuador.
Samples were collected between August 26-Septem- The complementarity between the two sites is the
ber 15, 1987 (Beni) and from September27-October proportion ofthe unique species to the pooled rich-
5, 1987 (Pakitza). Adult moths were collected by UV ness, or
light traps, spread, identified to family, sorted to mor-
U
pho-species, and counted. Voucher specimens have jk
been deposited in the National Museum of Natural (3)
History, Smithsonian Institution, Washington, D.C.
Collecting effort was defined as the number of trap Hemiceras species richness in westernAmazo-
nights: 12 and 8 trap nights in Pakitza and Beni, re- nia. The genus Hemiceras (Lepidoptera: Notodonti-
spectively. Otherstudies have includedthe numberof dae), representing 245 species, was used as an indica-
person-hours spentcollecting (Coddingtonetal. 1991, tor group for estimating species richness at three
Robbins et al. 1996), numberofcollecting days (Lou- rainforest sites: Pakitza and Rio Tambopata Research
tonetal. 1996), ortrapnights (approximately12hours Station in southeastern Peru, and Onkone Gare, in
inlength). Ecuador.
The Lepidoptera faunas ofPakitza and Beniwere Rio Tambopata Research Station, at 14°14'S,
compared using complementarity (Colwell and Cod- 69°11'W, is located on the Rio Tambopata, 30 air km
dington 1994). In comparing two sites, j and k, the SE of Puerto Maldonado, Madre de Dios, Peru, at
first site has a species richness of S. and the second 290melevation. Onkone Gare, at00°38'S, 76°36'W,is
site has Sk Ifthe number ofspecies in common be- aresearchstationwithinthe ReservaEthnicaWaorani,
tween both sites is V.k, then the total species richness Ecuador, at220 m elevation. Tambopatahadatotal of
forboth sites is 29 trap nights from November 2-25, 1979 and Sep-
tember 16-21, 1990. The 11 trap nights at Onkone
s.j,k =s.j + s.k-v.,jk,' (1) GarewereJanuary10, 12, 13-18,25;June20; andJuly
16, 1994.
andthe numberofspecies uniquetobothsites (U.k) is Species accumulation curves (Fig. 1) were used to
plot the cumulative number ofnew species collected
Ujk = Sj + Sk-2V]k. (2) overunit effort (number oftrap nights) at each ofthe
Volume 53, Number2 67
Fig.2. LocalitiesofCostaRicansitesusedinclusteranalysisofdissimilaritymatrixinTable3.
Fig.3. LocalitiesofSouthAmericansitesusedinclusteranalysesofdissimilaritymatricesinTables4and5.
three sites. To extrapolate total species richness from jackknife estimator ofspecies richness is based on the
the species accumulation curves at each site, four dif- numberofspecies thatoccurin onlyone sample L,
ferent nonparametric equations were compared: 1)
Chao 1, 2) Chao2, 3) first-orderjackknife, and4) sec- 1jackknife = S + L(n- 1/n), (6)
obs
ond-orderjackknife.
Chao (1984) developed a simple estimator of the where n is the numberofsamples.
true number of species at a given site based on the The second-orderjackknife (Burnham & Overton
numberofrare species in the pooled samplej. This is 1978, 1979) is liketheChao2estimatorwhere Listhe
consideredanabundance-basedestimatorbecauseitis number ofspecies that occurin only one sample and
M
based on the number ofspecies that are only repre- is the number ofspecies that occur in exactly two
sented by only 1 or 2 individuals to estimate overall samples,
species richness. Colwell and Coddington (1994)
calledthis Chao 1, 2jackknife
Sobs + V2 [L(2n-3)/n- M(n-2)2/n(n- 1)],
Chao 1 = So.bs+ a2/2b, (4) (7)
where S isthe observednumberofspeciesinasam- where nis the numberofsamples.
obs
ple, a is the number of species that are only repre- Hemiceras faunal comparisons between three
sentedbyone specimen in the pooled sample (single- tropicalregions. SpecieslistsofHemiceraswerecom-
tons), and b is the number ofspecies represented by piledfromspecimensinthe NationalMuseumofNatu-
two specimens in the pooled sample (doubletons). ral History, Smithsonian Institution, Washington, D.C.,
This estimatorworks wellwhen the samples contain a and were used to examine faunal relationships among
large number ofrare species (Chao 1984), which fre- sites in Costa Rica, Brazil, and western Amazonia. In
quentlyoccurs when sampling diverse groups such as Costa Rica, sixsiteswere chosentoseehowaltitude af-
insects. fectedspeciescomposition. The siteschosenwereJuan
A related estimatoris Chao 2 (Colwell & Codding- Vinas (1500 m), Tuis (732 m), Turrialba (634 m),
ton 1994),whichisbasedontheincidenceofrare spe- Guapiles (259 m), La Selva (40 m) and Sixaola River(0
cies amongsamples, m) (Fig. 2). Six sites in Brazil were chosen to examine
theeffectofforesttype: thelowlandAmazoniaforestin-
Chao 2 = S, +L2/2M, (5) cluded the sites ofSao Paulo de Olivenca (Amazonas)
obs
and PortoVelho (Rondonia), andthe Atlantic coast for-
where L is the number of species that occur in only est sites were Baixo Guandu (Espirito Santo), Campo
M
one sample, and isthenumberofspeciesthatoccur Bello (Rio de Janeiro), Hansa Humboldt (Santa Cata-
in exactlytwo samples. rina), and Santa Catherines (Santa Catarina) (Fig. 3,
Jackknife estimators (Burnham & Overton 1978, sites 1-6). Sixsiteswerechosentodeterminefaunalre-
1979) also use the distribution ofspecies among sam- lationships in western Amazonia. They included
ples (Colwell & Coddington 1994). The first-order Neblina,Venezuela; Onkone Gare, Ecuador; Sao Paulo
68 Journalofthe Lepidopterists' Society
Table1. Comparisonof%complementarityandnumberofspe- Table2. Estimatedtotalspecies richnessofHemicerasspecies
ciesofLepidopterabetweenBeni,BoliviaandPakitza,Peru. for4nonparametricestimators.
Family %Complementarity Numbersharedspecies Speciesrichness Pakitza Tambopata OnkoneGare
estimator Peru Peru Ecuador
Microlepidoptera
Cosmopterygidae 100.0 50(0) Observed 36 24 22
Tineidae 100.0 113(0) Chao1 43 29 40
Gelechiidae 98.8 167(2) Chao2 39 25 24
Oecophoridae 97.8 324(7) 1Jackknife 41 24 24
Pyralidae/Crambidae 96.8 281 (9) 2Jackknife 39 25 23
Macrolepidoptera
Noctuidae 98.6 296(4)
Notodontidae 95.2 63(3)
Geometridae 93.3 213(14) closer to the observed values than did Chao 1, on
Arctiidae 91.5 189(16) abundance-basedestimates (Table2).
Hemiceras faunal comparisons between three
tropical regions. Costa Bica: The high altitude sites
de Olivenca and PortoVelho in Brazil; and Pakitzaand ofJuanVinas andTuis (1500 m and 732 m) were clus-
TambopatainPeru (Fig. 3, sites 1-2, 7-10). tered at a dissimilarityvalue of0.389, and the lowalti-
Complementarity was calculated between sites. tude sites (0m and40m) ofSLxaolaBiverandLaSelva
Thesevalueswere usedto produce dissimilaritymatri- were clustered at a dissimilarity value of 0.571.
ceswhichwereconverted,foreasierinterpretation,into Guapiles (259 m) was clusteredwith the low sites at a
dendrograms usingclusteranalysis (SYSTAT 1992). dissimilarityvalue of0.667. The high altitude and low
altitude sites were clustered at a dissimilarityvalue of
Results
0.718. Turrialba (634 m) showed the greatest faunal
LepidopteracomplementarityinSWAmazonia. dissimilarity between all other sites at 0.758 (Table 3;
Some 38 families ofLepidoptera were collected from Fig. 4).
both sites, although onlynine families had numbers of Brazil: The two Amazonian sites of Sao Paulo de
speciessufficienttoillustratetrendsincomplementarity. Olivenca and Porto Velhowere clustered at a dissimi-
Five ofthese families were Microlepidoptera and the larity value of 0.883. Within the Atlantic Coast sites
remaining fourwere Macrolepidoptera. A total of1748 HansaHumboldtand SantaCatherineswere leastdis-
specimens representing 933 species were collected at similar (0.600), Campo Bello was most similar to
Beni and 1731 specimens representing 1006 species Hansa Humboldt + Santa Catherines (0.724), and
from were collected at Pakitza. The pooled species Baixo Guanduwas most dissimilarto the previous At-
richness forboth sites (S.k) was 1879, the total number lanticCoastForestsites (0.833). Dissimilaritybetween
ofunique species (U. was 1819, resulting in a com- the Amazonia and Atlantic Coast Forests was 0.951
k)
plementarityof96.8%. The Microlepidopterafamilies (Table 4; Fig. 5).
hadhighercomplementarityvalues (100-96.8%),than Western Amazonia: Pakitza and Tambopata in
the Macrolepidopterafamilies (95.2-91.5%), with the southeastern Peru were clustered at a dissimilarity
exceptionofthe Noctuidae (98.6%) (Table 1). value of0.766. Neblina, Venezuela and Onkone Gare,
Hemiceras species richness inwesternAmazo- Ecuador were clustered at a dissimilarity value of
nia. Comparingthe four nonparametric equations for 0.793. Thesefoursiteswereclusteredatadissimilarity
estimating species richness of Hemiceras at Pakitza valueof0.806. The faunaofSaoPaulode Olivengahad
and Tambopata, Peru, and at Onkone Gare, Ecuador adissimilarityvalue of0.815comparedtotheprevious
resulted in Chao 1 estimating the highest number of foursites andPortoVelhos faunawasthe mostdissim-
species. The incidence-based estimators (Chao 2 and ilar(0.833) (Table5; Fig. 6).
the Jackknifes) consistently estimated total richness
Table4. DissimilaritymatrixofcomparisonofBrazilianAmazo-
Table3. DissimilaritymatrixofCostaRicanaltitudmalsites. niaandAtlanticCoastForest.
JuanVinas SaoPaulode Porto Baixo Campo St.
(1500m) Tuis Turrialba Guapiles LaSelva Olivenca Velho Guandu Bello Catherines
Tuis(732m) 0.389 PortoVelho 0.883
Turrialba(634m) 0.771 0.758 BaixoGuandu 0.982 0.951
Guapiles(259m) 0.897 0.829 0.960 CampoBello 0.927 0.930 0.833
LaSelva(40m) 0.718 0.730 0.897 0.769 St.Catherines 0.945 1.000 0.867 0.724
SixaolaRiver(0m) 0.794 0.781 0.909 0.667 0.571 HansaHumboldt 0.958 1.000 0.870 0.792 0.600
) I
Volume 53, Number2 69
Turrialba634m (1)-
Tuis732m (2)
Juan Vinas 1500m(3)-
LaSelva40m (4)
SixolaRiver m (5)
Guapiles259m (6)
0.667 0.758
0.389 0.571 0.718
Fig.4. Dendrogramforclusteranalysisofdissimilaritymatrix(Table3)ofCostaRicansites.NumbersrefertolocalitiesonFig.2.Altitudes
areindicatedforeachsite.Thescaleindicatesdissimilarityvalue.
Discussion suitable richness estimator) shouldreachanasymptote
orremainconstantovertimewithadditionalsampling.
Lepidoptera complementarity in SW Amazo- Of the three sites, the species accumulation curve
nia. The lower complementarity found in most reached an asymptote, only at Pakitza (Fig. 1), sug-
Macrolepidoptera, comparedtothe Microlepidoptera, gesting that the species estimate there should be the
may be due to sampling bias; because the collecting most accurate. Although Tambopata is known for the
was done by UV light, larger moths may be coming high species richness ofvarious insect groups (Fisher
&
from a larger collecting universe than the smaller 1985, Paulson 1985, Pearson 1985, Wilkerson
moths, because the the larger species are better able Fairchild 1985, Robbins et al. 1996). The Hemiceras
to disperse than the smaller ones. The relativelyhigh fauna there is poor, considering that there are 245
complementarity of the Noctuidae is curious given species in the neotropics. Although samples taken at
that they are generally strong fliers, medium to large Tambopata and Onkone Garewere insufficient to ac-
moths, and many species are known for their migra- curately estimate richness, as shown by the non-
tion and wide ranging dispersal abilities. The answer asymptoticspecies accumulationcurves (Fig. 1), itap-
maylie in theirdiversity; because these moths are the pears from all the nonparametric estimates, and the
most speciose lepidopteran family, the sampling time steeperaccumulationcurve, thatPakitzaisevenricher
possible in this preliminary study may be inadequate inHemiceras thanisTambopata. Thatcontrastswith a
to accurately assess the ranges of many noctuid spe- study of the faunal relationships between the Ci-
cies, resultingin ahighercomplementarityvalue. cadoidea (Homoptera) (Pogue 1996) and Odonata
Hemiceras species richness inwesternAmazo- (Louton et al. 1996) at Pakitza and Tambopata, in
nia. Toaccuratelyestimate thetotalnumberofspecies which the species richness was greater at Tambopata.
of a target taxon (such as Hemiceras) at a particular Thus, eitherthepreliminaryestimateforHemiceras at
site the species accumulation curve (or the curve ofa Tambopata is inaccurate, or this study demonstrates
SaoPaulodeOlivenca(1
PortoVelho (2)
BaixoGuandu (3)
CampoBello(4)
St.Catherines(5)
5
HansaHumboldt(6)
0.883
-f— h
0.600 0.724 0.833 0.951
Fig.5. Dendrogramforclusteranalysisofdissimilaritymatrix(Table4)ofBrazilianAmazonianandAtlanticCoastForestsites. Numbers
refertolocalitiesonFig.3.Thescaleindicatesdissimilarityvalue.
—
)
70 Journalofthe Lepidopterists' Society
PortoVelho(2)
Sao PaulodeOlivenpa(1
Pakitza(7)
Tambopata(8)
Neblina(9)
OnkoneGare(IO)
0.806 0.833
H
I I I I
0.766 0.7930.815
Fig.6. Dendrogramforclusteranalysisofdissimilaritymatrix(Table5)ofwesternAmazoniansites. NumbersrefertolocalitiesonFig.3.
Thescaleindicatesdissimilarityvalue.
that different taxa, with divergent biologies, have dif- taxon can come from collecting, or from using mu-
ferent centers ofdiversity. seum collections to obtain faunistic data from specific
At the Onkone Gare site in Ecuador it is not clear sites. An advantage ofusing museum collections are
whether the species accumulation curve is approach- thedataavailablefrom sitesthatare nolongerpristine,
ingan asymptote (Fig. 1) andthere are more rare spe- such as those in Amazonia that were collected more
cies (12 singletons) than at the other sites, so the pre- than 50 to 100 years ago. Today, with the destruction
diction of 40 species by Chao 1 (Table 3) may be ofthe rain forest, these sights will no longerhave the
accurate. Based on the species richness data of same biota. The target taxon has to be common
Hemiceras, thefollowingare recommendedforfollow- throughout the study area so it can be easilysampled
up studies: 1) collecting effort must be adequate for andthere shouldnotbe adominance ofrare species.
the species accumulationcurve, orthe estimate curve, Hemiceras faunal comparisons between three
to reach an asymptote, 2) ifthere is apreponderance tropical regions. The six sites in Costa Rica show a
of rare species, (singletons) Chao 1 should give the broad altitudinal range from 0-1500 meters. Altitude
highest, andperhaps bestestimate, and3) ifthe num- seems toinfluence species composition amongsites in
ber ofrare species is low, Chao 2 orthe second-order Costa Rica more than distance between sites. Juan
jackknife may give a better estimate. It is also impor- Vinas (1500 m) andTuis (732 m) are the highest sites
tantto chooseyourtargettaxon carefully. and are clustered at a dissimilarityvalue of0.389, the
To assess taxon richness, targettaxacanbe anycat- lowest ofanypair. Ifdistancewas the significantlimit-
egory, an order, family, subfamily, tribe, orgenus. The ingfactoroffaunalcomposition, onewouldexpectthat
taxon should be chosen with care. For example, one Tuis and Turrialba would be clustered. The same is
that is too speciose requires too much time to process true for the lower altitudinal sites, with La Selva (40
andextrapolatetheneededdata, onewithtoofewspe- m) and Sixaola River (0 m) having the lower dissimi-
cies could result in insufficient data. I have found in larityvalue (0.571), even though La Selva is closer to
the Neotropics that atarget taxon of200-400 species Guapiles than the Sixaola Riversite (Table 3) (Fig. 4).
seems to be large enough so that the species accumu- Analysis ofthe six sites within Brazil were used to
lation curve reached an asymptote after 20-30 sam- showiftherewas afaunaldifference betweenAmazo-
ples (trap nights, in this case). The dataforthe target nia and the Atlantic Coast forest. The Atlantic Coast
sites were clustered (Baixo Guandu to Hansa Hum-
boldt) and were quite distinct from the Amazonian
fauna (Sao Paulo de Olivenca and Porto Velho) which
Table5. DissimilaritymatrixofAmazoniansites.
were also clustered. Within the Atlantic Coast Forest
Onkone SaoPaulode Porto distance seems to be influencing the faunal relation-
Neblina Gare Olivenca Velho Pakitza
ships. Hansa Humboldt and Santa Catherines were
OnkoneGare 0.793
S.P. DeOlivenca 0.918 0.873 most faunistically similar and closest in distance.
PortoVelho 0.947 0.886 0.883 Campo Belloshows similaritywith HansaHumboldt +
Pakitza 0.860 0.863 0.831 0.833 Santa Catherines and is closer than Baixo Guandu,
Tambopata 0.806 0.821 0.815 0.915 0.766 which is the most dissimilar and furthest from these
Volume 53, Number 2 71
ntawlodsiitfefs.erCelnucsetebreatnwaelyesnisAimnadizcoanteisathaantdthtehreeAistlaanftaiu-c Coddnianrgatnodna,&J.S.A.F,.CL.arEc.heGrr.is1w9o9l1.d,DeDs.igSniilnvgaaDnadvitleas,tinEg.saPme--
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