ConselVation value of light trap-collected caddisfly assemblages(Insecta: Trichoptera) was evaluated on the basis of their species richness. The assemblage coming from an artificial stream showed a higher conservation value based on species richness than the natural ones. In contrast, using rarefaction, the conservation value of the assemblage in the artificial stream was lower in comparison with assemblages in natural habitats. Further examples are given to demonstrate the importance of rarefaction in comparing species richness of assemblages

Schmera, Dénes

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Opusc.Zool. Budapest, 2002,34: 77-83Comparison of species richness of light trap-collectedcaddisfly assemblages (Insecta: Trichoptera) usingrarefactionD. SCHMERA*Abstract. ConselVation value of light trap-collected caddisfly assembIages (InsectB.: Trichoptera) was evaluated onthe basis of their species richness. The assemblage coming from an artificial stream showed a higher conservation valuebased on species richness than the natural ones. In contrast, using rarefaction, the conservation value of the assem-blage in the artificial stream was Iower in comparison with assemblages in natural habitats. Further exampIes are givento demonstrate the importance of rarefaction in comparing species richness of assemblages.H uman environmenta! disturbance and its ef-fect on biota are one of the most importantphenomena for community ecologists (pianka,1970; Southwood, 1977). Falling biodiversity (Ju-hász-Nagy, 1993) has been the fIrst and mostsignificant sign calling cur attention to changes instructure in the biosphere. This process is ofglobal scale, however, could also be demonstratedby local studies. For instance, butterfiy com-munities at the foot of Fuji Mountains (Japan)show sensitivity (reduction in species richness) tohuman disturbance (Kitahara & Fujü, 1994; Kita-hara & Sei, 2001). Nowadays, it is generally ac-cepted that degradation processes could be meas-ured through varlous community structural char-acteristics. Among others, species richness is thesimplest measurement to indicate degradation.Generally, a wide array of species represents a.wen being" state of community (high conservationvalue) , while law species richness does not (lowconservation value) (Magurran, 1988). This con-ception is strongly supported in aquatic ecology,where biotic indices (for instance Ephemeroptera-Plecoptera-Trichoptera index) are chiefly based onspecies richness (Stone & Wallace, 1998). Accord-ingly, numerous studies have been conductedusing species richness as a measure of environ-menta! conditions (Ivol & al., 1997; DeWalt & al.,1999; Ruse & Herrmann, 2000; Lomond & Colbo,2000). In addition, species richness is also com-monly used in measuring seasonality of phy-toplankton (Padisák, 1993) or in comparing mac-roinvertberate assemblages in streams (Schmera,1999; Andrikovics & Kiss, 2000; Csörgits, 2000;Kiss & al., 2001) or carabid communities (Magura& Tóthmérész, 1996). The number of species in asampie would be a proper measure of the speciesrichness of a studied community, however, wescarcely are in the position to collect alI the or-ganisms (species and individuals) in the givencommunity. Based on tieId observations (Gotelli &Graves, 1996), the more individuals are sampled,the species richness rises unti! an asymptote isreached. Several hypothesises have been proposedto expIain the phenomenon. The most commonlyknown hypothesis include a passive sampling mo-del, where richness is Iarger because of thestatisticalIy greater probably of sampling newspecies in a large sampIe (Giller & Malmqvist,1998). Consequently, species richness yieldingfrom samples significantly differing in sizes(number of specimens) could not be compared.Therefore, the general purpose of this studywas to demonstrate the advantages using rare-faction in community comparisons based onspecies richness. Rarefaction is a mathematicalmethod, thereby the species richness of com-munities could be compared, as if their number ofindividuals where the same. This method has beenused in comparing carabid communities (Magura& Tóthmérész, 1996) or macroinvertebrate assem-blages (McCabe & Gotelli, 2000). As species rich-ness of stream dwelling caddisflies is strongiy de-.Dénes Schmera, MTA Növényvédelmi Kutatóintézete (pJant Protection Institute of the Hungarian Academy ofSciences), 1022 Budapest, Herman Ottó út 15, Hungary.Schmera: Species richness of caddisfly assemblagesTable 1. Comparison of sampling circumstancespendent on the stream order (Wiberg-Larsen & al.,2000) and size (Vilson & Hawkins, 1998), thereforeassembiages representing streams and rivers wereseparately studied. Specifically, caddisfly (Insecta:Trichoptera) assemblages collected by light trapson the bank of streams and rivers were separatelycompared using the collected by light traps on thebank of streams and rivers were separatelycompared using the collected number of speciesand the rarefaction of each caddisfly assemblagesfrom own and a literature data set.sosaprobic to alpha-mesosaprobic, Chantara-mongkol, 1983).The identification of Hydropsyche females iscurrently not possible to species level (Malicky,1983), so they are used as a new _species- in theanalysis. Consequently, species richness could beincreased (8+1) comparing with one in the puh-lished data.The rarefaction solutionMATERIAL AND ME'nIODSOrigin of the dataSix caddisfly assemblages of streams and nyerswere collected by light traps installed at the bankof the waters (Table 1). Assemblages A and Cwerecollected at an undisturbed, nature reaches of thestreams, while assemblage B sprang from anartificial stream «<Meleg-víz-; Uherkovich & Nóg-rádi, 1999). «Meleg-víz- canies the warm karsticwater of a bauxite mine. Thus, in February, itstemperature does not falI below 170C even ap-proximately 15 km away from the pumping station(Uherkovich & Nógrádi, 1999). Rivers were re-presented by assembiages D, E and F. Assem-blages D and F carne from the River Danube, whileassemblage E from the River Tisza. Danube, asother large rivers in Europe, is heavily affected bydomestic and industrial sewage, therefore itswater quality can be classified as 2-3 (betame-The number of species shows increasing func-tion as the number of collected individuals growth.To solve this phenomenon, rarefaction intends toevaluate the number of species of a theoreticalassemblage defined by its number of individuals.Consequently, assemblages differing in number ofindividuals can be compared in a way, as if theirnumber of individuals were the game. Therarefaction could be calculated on the basis of aprobabiy calculation or randomisation. The ex-pected number of species of a theoretical com-munity could be evaluated on the basis of thetheoretical number of individuals and on the basisof the relative abundance vectors of the realisticcommunity (Hulbert, 1971; Tóthméresz, 1995):srES(m) = ST -L(l-pi)m;=1where ES is the expected number of species at mtheoretical number of individuals, while pi is therelative abundance of the í-th species in therealistic community characterised by ST species.78Schmera.' Species richness of caddisjly assemblages79~ 10 100 1000 10000 100000 CNFigure 1. The comparison of species richness of stream assemblages (A, B and C) using rarefactionTable 2. species richness and number ofÚ1dívidua/s of the studíed assemblagesAssem blage Species richness Number of individualsA 53 8065B 62 90506c 34 1395D 26 831E 27 6164F 48 22882RFSULTSSpecies richness of stream dwelling caddisflyassembJagesAmong the stream assembiages, the speciesrichness of assembiage B is the highest followedby A and C (Table 2). Based on the measurementof species richness, the assemblage of an artificialstream represents a high conselVation status,while the nature ones do not. Stream dwellingcaddisfly assemblages showed the following rankof species richness obtained on the basis ofrarefaction measured through probably: assem-blage A had the highest estimated species rich-ness followed by C and B (Fig. 1). Using rarefac-tion with randomisation, significant difference wasfound in stream assembiages between speciesrichness of assembiage A and assembiage B andbetween assemblage B and C, respectively (Table3). No significant difference was found between thespecies richness of assembiage A and C (Table 3).At each comparison, ES was calculated be-tween m =10 to min(Nj), where Nj represents thenumber of individuals of j assemblage (Table 1).ES(m) was calculated using the DIVORD computerprogram (Tóthmérész, 1993, 1994).Unfortunately, such a kind of evaluation ofspecies richness makes it impossible the statis-tical comparison of difIerent assemblages, as onlythe expected number of species with the highestprobably is given. By comparison of two as-semblages using randomisation, the assemblage(x) represented by higher number of individualswas rarefied to the abundance level (number ofindividuals, aj of the smallest assemblage (y) 1000times in the following way: a individuals wererandomly sampled from the assemblage x (Gotelli& Graves, 1996). In each case, the number ofspecies will also be obtained. Thereby, mean andvariance of the expected number of species ofassemblage x could be obtained at the abundancelevel of the assemblage y. The difIerence betweenthe two values could be calculated on the basis ofthe normaI distribution (Gotelli & Entsminger,2001). Randomisation and the mean and varlanceof the expected number of species were caIculatedusing the ECOSIM computer program (Gotelli &Entsminger, 2001). The difIerences between thetwo values were calculated on the basis of theprobably distribution calculator of STATISTICAcomputer program (StatSoft 2000).Species riclmess of river dwelling caddisflyassembJagesComparing species richness of assemblagescarne of the rivers, assemblage F is the mostvaluable one followed by E and D, respectivelyífable 2). However, caddisfly assemblages comingfrom the rivers show the following rank after rare-80Schmera: Species richness of azddisjly assemblagesTable 3. Expected number of species of stream assemblages (observed value ís given as mean::!: variance,NS denote non significant dífference, *** means highly significant (p<0. 001) dífference}Assem blage Rarefied to the abundance level ofassembiage ARarefied to the abundance level ofassembiage Cobserved value signit1cance observed value significance37.6 :t 5.118.4:t 4.9NSA32.8 ::t: 7.8.-B..Table 4. Expected number of species of river assemblages (obseroed value is given as mean ::t va1Íanre,NS denote non.significant difference, *** means highly significant (p<O. 001) difference}AssembIage Rarefied to the abundance level ofassemblage DRarefied to the abundance level ofassemblage EobseIVed value signitlcanceobserved value significance...14.3:1: 3.49.9:1:4.1E27.5:t7.3F .-faction: assemblage D has the highest estimatedspecies richness at alt m values followed by E andF (Fig. 2). The curve of E and F cross each other,therefore the rank of the estimated species rich-ness depends upon the abundance level (numberof individuals) , at which the comparison wouldmade. In river assemblages (Table 4), significantdifference was found between assembtages D andE and between D and F. No significant differencewas found between assemblages E and F.DISCUSSIONdomisations could not conf1rm the obtained rankas no significant difference was found between thespecies richness of assemblage A and C. Bycomparing assemblages representing rivers,species richness indicated the reversed order thanon the basis of rarefaction (F>E>D vs. D>E---Fj.Consequently, if conselVation value would bemeasured through the number of species ornumber of expected species, different solutionscould be obtained. For instance, even thoughspecies richness indicated the highest conselVa-tion value of an assembiage coming from anartificial stream, rarefaction rejected this assump-tion indicating another rank of conselVation value.Obviously, the differences between the twoapproaches (species richness and rarefaction so-lutions) carne chiefiy from the differences innumber of individuals (sarnpling effort, Table 2)and from the distrIbution of individuals arnongspecies (abundance). For instance, white assem-biage B was represented by 90,506 individuals,assemblage C by only 1,395. While at a small as-sembiage size, the presence of a rare species wassmall, in a big one it was higher. Therefore, aSpecies richness is the simplest way todescribe community or regional diversity (Magur-ran, 1988). Species richness shows sensitivity tohuman activity (Kitahara & Fujü, 1994; Kitahara& Sei, 2001), therefore, it can be used as ameasure of conservation value.The comparison of species richness in streamassemblages showed the B>A>C rank, while onthe basis of rarefaction using probably calcula-tions A>C>B. However, rarefaction based on ran-81Schmera: Species richness of caddisjly assemblagesspecific species-individual curve could be ob-tained: number of species grows as the number ofcollected species increases (Magurran, 1988). Theconcept of rarefaction was efficientiy used in com-parison of species richness of stream macroinver-tebrates (McCabe & Gotelli, 2000) or in compar-isDn of number of passerlne hird species at dif-ferent territorial pairs (Gjerde & Saetersdal, 1997).In addition, Gjerde & Saetersdal (1997) noted thatdiversity indices, like Shannon (H1 or Símpson'sindex (D), could not always be used in comparisonof diversity of communities.In spite of its ímportance, ecologists do notalways consider the effects of abundance andsampling effort on species richness measures andcomparisons. Overall, rarefaction solution allowsfor meaningful standardisation and comparison ofdatasets (Gotelli & Colwell, 2001). This study de-monstrated that using rarefaction in comparisonof assembiages with great differences in number ofindividuals was recommended.Acknowledge~nt8. 1 thank Dr. Arpád Szen-tesi and Prof. Péter Biró for their helpfui com-ments on the manuscript.REFERENCES(EPr) species richness and commUIÚty structure inthe Lower Illinois River Basin of illinois. The GreatLakes Entomol., 32: 115-132.GJERDE, 1. &. SAETERSDAL, M. (1997): Effects of aviandiversity of introduction spruce Picea spp. planta-tions in the native pine Pinus sylvestris forests ofWestern Norway. Biol. Conseroatíon, 79: 241-250.GoTELU, N. J. &. COWELL, R. K. (2001): Qualifying bio-diversity: procedufeS and pitíalls in the measurementand comparison of species richness. Ecol. Letters, 4:379-391.GoTELU, N. J. &. ENTSMINGER, G. L. (2001): EcoSim: Nullmodels software for ecology. Version 7.0. AcquiredIntelligence Inc. &. Kesey-Bear. http://homepages.together .net/-gentsmin/ ecosim.htm.GoTELLI, N. J. &. GRAVES, G. R. (1996): Null models inecology. Smithsonian Inst. Press, Washington DC.HULBERT, S. H. (1971): The nonconcept of species di-versity: a critique and alternative parameters.Ecoiogy, 52: 577-585.IVOL, J.-M., GIUNAND, B., RICHOUX, P. &. TACHET, H.(1997): Longitudinal changes in Trichoptera andColeoptera assemblages and environmental condi-tions in the Loire River (France). Arch Hydrobiol.,138: 525-557.JUHAsz-NAGY, P. (1993): The fading diversity. (InHungarian.) sdencia Kiadó, BudapestKIss O., ANDRIKOVICS, S., MuRANYI, D. &. LIP!'ócZY, Á.(2001): The aquatic insect fauna in the CsörgõStream of the Máu-a Mountains (Hungaxy). (InHungarian.) HidroL Kózlóny, 81: 392-393.KITAHARA, N. &. FuJII, K. (1994): Biodiversity andcommUIÚty structure of temperate butterlly specieswithin a gradient of human disturbance: an analysisbased on the concept of generalist vs. specialistsu-ategies. ReseaTcoJ1 in Population Ecology, 36: 187-199.KITAHARA, N. &. SEI, K. (2001): A comparison of diversityand structure of butterfiy communities in semi-natW'al and human-modified grassland habitats atthe foad of Mt. l'i.lji, central Japan. Bíodíversíty &.ConseTllatíon, 10: 331-351.LoMOND, T. M. & COLBO, M. H. (2000): Variations inlake-outlet Ephemeroptera. Plecoptera, and Tricho-ptera communities amongst regions of eastern New-foundland, Canada. Canad. Joum. Zool., 78: 1536-1543.ANORIKOVICS, 8. & KIss, O. (2000): Bioindicatioo by mac-roinvertebmtes in tlle Danube. 3. Iovestigatioo 00tlle aquatic insect laIVae in tlle Hungarian 8ectioo ofDanube. Hidra!. K6zl6ny, 80: 272-274. (In Hun-garian.)ANDRIKOVICS, 8., KIss, O., NOSEK, J. & OERTEL, N. (2001):Bioindicatloo by macroinvertebmtes in tlle Danube.7. PossibiIity of light-trapping for the study of aquaticmacroinvertebmtes. Hidra!. K6zlöny, 81: 314-315. (InHungarian.)CHANTARAMONGKOL, P. (1983): Light-tmpped caddistlies(rrichoptem) as water quality indicators in largerivers: Results from tlle Danube at Veroce, Hungary.Aquatic Insects, 5: 33-37.CSóRGITS, G. (2000): Comparisoo of long-term changes ofinvertebmte macrofauna in two streams of tlle PilisMountains, Hungary. Opusc. Zoo!. Budapest, 32: 27-49.DEWALT, R. E., WEBB, D. W. &. HARRIS, M. A. (1999):Summer Ephemeroptera, Plecoptera, and Trichoptem MAOURA. T. &. TÓTHMgRgSZ. B. (1996): Comparison of the82Scfunera: Species richness of caddisfly assemblagesNorth Hungary. Opusc. Zool. Budapest, 33: 83-89.SOUTHWOOD, T. R. E. (1977): Habitat, the templet forecological strategies? Joum. Animai Ecol., 46: 337-365.StatSoft, Inc. (2000): STATISTICA for Windows (Com-puter program manual). 7Ulsa, OK.ToNE, M. K. &. WALLACE, J. B. (1998): Long-term recoveryof a mountains stream ftom cleaI1:ut logging: theeffects of forest succession on benthic invertebratecommunity structure. Freshwater Biology, 39: 151-169.TóTHMtRtsZ, B. (1993): DivOrd 1.50: A program for di-versity ordering. Tiscia, 27: 33-44.TarHMtRtsZ, B. (1994): DivOrd 1.60: Diversity ordering:finite and infinite samples. 7Yscia, 28: 63-65.TóTHMERtsz, B. (1995): Comparison ofdifferent methodsfor diversity ordering. JOUnL Vegetation Scie~, 6:283-290.carabid communities of a ronal Queroeto-c.arpinetumand pine pJantations. proc. "Research, Consenlation,Management" Conl Aggtelek, Hungary, p. 131-135.MAauRRAN, A. E. (1988): Ecological diversity and itsmeasurement. Croom Heim Ltd., London.MALICKY, H. (1983): Atlas of European Trichoptera. Junk,The Hague.MCCABE, D. J. & GOTELLI, N. J. (2000): Effects ofdisturbance frequency, intensity, and area on assem-blages of stream macroinvertebrates. Oeoología, 124:270-279.PADIsAK, J. (1993): The influence of ditTerent disturbancefrequencies on the species ricJ:mess, diversity andequitability of phytoplankt.on in shallow lakes. Hyd-robiologia, 249: 135-156.PlANKA, E. R. (1970): On r- and K-selection. AmericanNaturalist, 102: 592-597.RUSE, L. P. & HERRMANN, S. J. (2000): Plecoptera andTrichoptera species distribution related to en-vironmental charactersitics of the metal-polluted Ar-kansas River, Colorado. Western N. Amerioon Na-turalist, 60: 57-65.SCHMERA, D. (1999): Change of structural charactersticsof caddisflies (Insecta: Trichoptera) along the Ber-necei Stream (Börzsöny Mountains, Northern Hun-gary). (In Hungarian.) Tennészetvéd. K6zlem, 8: 173-183.UHERKOVICH, A. & NOORADI, S. (1999): Caddistlies rrri-choptera) of artificial water courses in the BakonyMountains, Central Hungary o Braueria, 26: 21-23.VILSON, M. R. & HAWKlNS, C. P. (1998): Biodiversity ofstream insects: variation at local, basin and regionalscale. Ann. Rev. Entomol., 43: 271-293.WIBERG-LARSEN, P., BRODERSEN, K. po, BRICKHOLM, So,GRON, Po No & SKRIVER, J. (2000): Species richnessand assemblage structure of Trichoptera in Danishstreams. Freshwater Biology, 43: 633-6470SCHMERA, D. (2001): Seasonal activity of caddisflies(Insecta: Trichoptera) in the Börzsöny Mountains,83

A comparison of diversity and structure of butterfiy communities in seminatW'al and human-modified grassland habitats at the foad of Mt. l'i.lji, central Japan. Bíodíversíty &.

Atlas of European Trichoptera.

Biodiversity and commUIÚty structure of temperate butterlly species within a gradient of human disturbance: an analysis based on the concept of generalist vs. specialist su-ategies.

Biodiversity of stream insects: variation at local, basin and regional scale.

Bioindicatioo by macroinvertebmtes in tlle Danube. 3. Iovestigatioo 00 tlle aquatic insect laIVae in tlle Hungarian 8ectioo of Danube.

Caddistlies rrrichoptera) of artificial water courses in the Bakony Mountains, Central Hungary o

Change of structural characterstics of caddisflies (Insecta: Trichoptera) along the Bernecei Stream (Börzsöny Mountains,

Comparison ofdifferent methods for diversity ordering.

Comparisoo of long-term changes of invertebmte macrofauna in two streams of tlle Pilis Mountains,

DivOrd 1.50: A program for diversity ordering.

DivOrd 1.60: Diversity ordering: finite and infinite samples.

Ecological diversity and its measurement. Croom Heim Ltd.,

EcoSim: Null models software for ecology. Version 7.0. Acquired Intelligence Inc.

Effects of avian diversity of introduction spruce Picea spp. plantations in the native pine Pinus sylvestris forests of Western Norway.

Effects of disturbance frequency, intensity, and area on assemblages of stream macroinvertebrates.

Habitat, the templet for ecological strategies?

Long-term recovery of a mountains stream ftom cleaI1:ut logging: the effects of forest succession on benthic invertebrate community structure.

Longitudinal changes in Trichoptera and Coleoptera assemblages and environmental conditions in the Loire River (France).

Null models in ecology. Smithsonian Inst.

On r- and K-selection.

Plecoptera and Trichoptera species distribution related to environmental charactersitics of the metal-polluted Arkansas River,

PossibiIity of light-trapping for the study of aquatic macroinvertebmtes.

Qualifying biodiversity: procedufeS and pitíalls in the measurement and comparison of species richness.

Species richness and assemblage structure of Trichoptera in Danish streams.

STATISTICA for Windows (Computer program manual).

Summer Ephemeroptera, Plecoptera, and Trichoptem

The aquatic insect fauna in the Csörgõ Stream of the Máu-a Mountains (Hungaxy).

The fading diversity. (In Hungarian.) sdencia Kiadó,

The influence of ditTerent disturbance frequencies on the species ricJ:mess, diversity and equitability of phytoplankt.on in shallow lakes.

The nonconcept of species diversity: a critique and alternative parameters.

Variations in lake-outlet Ephemeroptera. Plecoptera, and Trichoptera communities amongst regions of eastern Newfoundland,

Comparison of species richness of light trap-collected caddisfly assemblages (Insecta: Trichoptera) using rarefaction

Comparison of species richness of light trap-collected caddisfly assemblages (Insecta: Trichoptera) using rarefaction

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