Spiders in Natural Pest Control: a Review

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Spiders in natural pest control: a review

Author(s): Nyffeler, Martin; Benz, Georg

Publication Date: 1987

Permanent Link: https://doi.org/10.3929/ethz-a-005802284

Originally published in: Journal of Applied Entomology 103(1-5), http://doi.org/10.1111/j.1439-0418.1987.tb00992.x

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ETH Library Departrnent of Enton'tology, Szoiss Federal Institute of Technology, ETH-Zentrum, Zaicb, Sntitzerland

Spiders in natural pest control: A reviewl

By M. Nvnnnrnn and G. BnNz

Abstract In spite of the large number of studies about the ecology of spiders carried out in the last two decades in different types of ecosysrems all over the world, the significance of these animals as natural controI agents is still largely unknown. In this paper the literature about that subject is reviewed. Totally 300 scientific papers, published betwein 1920 md 1984, are cited here. Several European and American studies have provided evidence. that in undisturbed grassland ecosystems and forest ecosystems spiders can play an important ecological role as preäators of inseits and other invertebrates. Alsd in orchards, Äo, ,.."äd with pestic"ides, that are'to a certain degree comparable wirh forest ecosystems, spiders can be abundant predators. In contrast to that, the opinion about the predatory importance of spiders inhabiting cultivated fietds is controuersial. The^results of some'Europ."n studies indicatd, that the foliaie-dwelling spid_ers of cultivated fields, because of their low pipulation densities, are of minor importance as predators of insects. Other European studies show,-that the ground-dwelling spiders oT cultivated fields are concerning their abunäance a dominant Dredator-group, those significance as control agents still is largel! unknown up to the p."..nt. In rice lieldi (swamp äcosystems) in Asia, receiving little or no pesticides, as well as in European and American swamp ecosystems, spiders may De an rmportant predator group. In houses in South Africa spideis were used successfully as biological control agents against flies.

1 Introduction

Spiders are among the most abundant predarors of insects of terrestrial ecosystems (Trscnrrn 1965; veu Hoor 1971;MovrDER and Pir,rc:r^rn 1972; ScnerrBn 1974; EowrRDs er aI. 1,976). Under favorable conditions they can reach maximal densities of up to 1000 individuals/m2 approximarely (Pnensr 1946; Durruy 1962; \ürrorueNN 1978). It has therefore been supposed for some time that spiders may play an important role as stabilizing agents and/or regulators of . insect populalions in agroecosystems, forest eöosystems, and other terrestrlal ecosystems, Qyri"g the_ last 2(i years,numerous studies on the spider faunas in agricultural habitats have been published all over the world. Tables 1-4 give a ieview classified according to cöntinents and countries: table 1 for Euroie (including the U.S.S.R.), table 2 for America, table 3 for Asia, and table 4 for Africa anil the Oceanic-Australian region. Also in forest ecosystems many studies on lThis paper represents a revised portion of a thesis submitted in 1982 by the first author to the Swiss Federal Institute of Technology (ETH) at Zurich in partial fulfillment of the requirements for the Ph.D. degree (Doctor of lrüt"..l Sciences). This iesearch was supported by'the Swiss National Science Foundation, grant 3.0020.76.

U.S. Copyright Clearance Center Code Sratement: 0044-2240/87/0304-0321, $ OZ.SOUO J. Appl. Ent. 103 (1987),321-339 @ 1987 Verlag Paul Parey, Hamburg und Berlin ISSN 0044-2240 322 M. Nyffeler and G. Benz

Table 1. List of literature on the spider faunas in agroecosystems in Europe, including the U.S.S.R.

Country Agroeco- Country Agroeco- Authors system system

Austria meadows Luuar (1979) German rape Brvrn (1981) Tnaren et al. (1977, Demo- sugar-beet BrrEn (1981) 1978) cratic kohlrabi Bnvrx (1981) arable Tnerrn and SrrrNpn Republic marrow- Bnvrx (1981) land (1e75) stem kale TruBn et al. (1977) Great pasture Cnrnnrrr (1964) orchards Püxnrxcrn (1982) Britain Dunrrv (1962) meadows Dumnv (1924) Belgium cereals Dr Clencq (1979) cereals Fnasen (1982) CortrNre md Crencq Locrur (1928) (1e77) VrcrpnrrraN and Surorn- diverse BosuaNs and Corrrnrr reuo (1975) crops (1e77) POtato DUNN (1949) sugar-beet Tuonr.rHrn (1983) Bulgaria pasture Drrcssv and KaJar orihards (1956) (re74) CmNr Hungary allalfa Barocn and Loxsa Czecho- pasture Pornltac (1968) (1e56) slovakia alfalta Mrrun (cit. Luczar, Noruay straw- Taxso,r.r (1973) 1979) berry sugar-beet Mnren (1974) Poland pasture Drr-cnrv md K-qer Federal meadows Boress (1953) (1e74) Republic Scnarrpx and Hers meadows BneYl,rEyen (1967, 197 8) of (1e7e) Kalar (1960, 1962, 1971, Gernany clover BoNrss (1958) 1978,1980) alfalfa Bourss (1958) Krler and Jarurczvr cereals Besroov (1973) (1e75) BesBoow md Mrrrrr Kayar et al. (1968,1971) (re77) alfalfa Lvcz* (1975) Bnassr (1975) cereals Lucz K (1974, 1975, asparagus DrNcren (1935) 1976,1979) vineyard Krnll (1978) POtato Czlyra and Kerre (1976) orchards Knauvrcn (1961) G.uecxe (1966) Naror (1976) Lucz^K (197+,1975, diverse Hevoru.rlw (1953) 1976, 1979) crops TrscHren (1958) sugar-beet Cza1x.r. and Goos (1976) Goos (1973) Finland leys Hunra and Sweden straw- Aruqursr (1981) R*rmrwrr (1924) berrY cereals Huxre and Rmrlrarrer (1924) Switzer- meadows Brrz & Nvrnrren (1980) land (1974,1975) Rut:rrunrl and Meunrn Huura (1968) Nvnrnrrn and Br.Nz (1979b,1981b) Frmce alfalfa CHauvrr (1960,1967) cereals Nnrrlen and BENz (1979a, 1980a) Germm meadows Bercn (1981) Nvnnrrrn md Belrz (1981a,1981b) Demo- Mürrrn et al. (1978) cratic grass- Brvrn (1981) Nvnrnrrn md Brrz Republic clover (1982a, r982b) mrxture rape Nyrrrrrn and BBNz alfalfa Gurrn (1963) (1979a) cereals Brrrn (1981) U.S.S.R. meadows Vrnesrr (1965) DrErnrcn and Görzr cereals AsHrnavrv (1923) (1e74) POtato Kovar (1975)

spiders have been carried out (review in VIrf 1953; KlncHnnn 1964; Mour- DER and RrrcHn 1972; Nyrnnmx 1982a). Furthermore, spiders have been studied in garden ecosystems (e.g. NvrrnrER 1983). In spite of the large number of existing studies, the significance of the spiders as natural control agents in agroecösystems, forest-ecosystems, and

l- Spiders in natural pest control 323

Table 2. List of literature on the spider faunas in agroecosytems in America

Country Agroeco- Authors Country Agroeco- Authors system system

Brazil sugarcane Bamosa et al. (1979) U.S.A. sweet corn Evrnrv (1938) rice $0ooos and Henn* over- (1966) Canada Tunxnurr (1e76) grazed pasture soybeans Brrcrsrslenrand HucceNs (1962) meadows Doroern (1971,1977) CurrN and Rusr (1980) Doroern and BrNrs al. (1976) 1977 Dnnz et Lrsen and DoNoen et al. lJxzrcrrr (1978a, b) (1970, 1972) TunNrnsrtn (1925) Fox and Donoerr (1972) (1980) wheat DoeNr and Doroerr Vurrcour (1e7e) cole Prunnrrr- (1961) orchards DoNoerr (1956, 1958) guar Rocrns and HonNrn Doroern et al. (1979) (1e77) Puruer (1967) sugarcane HrNsrrv et al. (1961) Purue.N and Hrnnr Nrcu md Hrnsrrv (1e66) (1e6e) cotton Bunrercn et al. (1973) Panama seed reser- Bmvurven (1928) Cram and Grrcr (1961) vation DraN et al. (1982) pasture (1928) Bnrvurvrn Donnrs (1970) banana HARRTSoN (1968) et al. (1976) plants JonNsow KeceN (1943) Peru cultivated Acurran (1965) Lrrcn and Hunllr fields (1e6e) cotton Acurren (1974) Locxrny et al. (1979) U.S.A. pasture Hoveno and Orrven McDeuer and SrrnrrNc (1e78) (te7e) Prcr and \(urcoun Prnruuen (1964) (1e78) Prerrns and Srrnrrrc '\üünrrcoun et al. (1963a) (1e74) Iüolcorr (1937) Snlpenn and SrrnrrNc meadows \Tolcorr (1937) (1e72) Srau et al. (1978) allalla Hovnrr and 'S7Hrrcovg Pronrovsrr (1971) and Bln (le64) Murrepper and Cxme 'rüünrrcoMs (1eZob) and Teorc Scnrrncnn and Dnrnrcr (le63) (1e60) \Tnrrcoun et al. Vurnrn (1923) 1963a, b) Yeancar.r (1975a, b) citrus CARRoLL (1980) Yr.q.nceN and CorHmr Muur (1973, 1975) (1e74) orchards Lpcrqen and OarueN Yrencax and Dotoere (le64) (|e74) McCerrney and cereals HonNrn (1972) HonssuncH (1978,1980) Murr.lppeN and CHme Specsr and Dotoau (t970a) (1e60) grain Beruv and Cseoe diverse BrrsrNc (1920) sorghum (1e68) crops $Tnrlcoun(1974,1975) other terrestrial ecosystems is still largely unknown. This could be attributed to the fact that up to the present most studies were limited either to the investigation of the species composition and the seasonal occurrence of spiders in the field (sweep net and pitfall trap collections) or to measurements of respiration, food consumption tests, and prey preference trials in the laboratory. However, the results of investigations on the nutrition of spiders in the laboratory cannot readily be extrapolated to field conditions, beCause spiders may behave differently in the laboratory than in the field. In the laborätorv, certain hunting spiders consume considerably more prey than would be 324 M. Nyffeler and G. Benz

Table 3. List of literature on the spider faunas in agroecosytems in Asia

Country Agroeco- Authors Country Agroeco- Authors svst€m system

India naize Snenue and Sanup Japan rice Tovooe and Yossruune (1e7e) (r966) SrncH and Saronu YecrNuue (1965) (1e76) YeueNo (1977) rice Karonr (1976) cabbage KevesHrue (1960) Salrar and Mrsna (1975) Suzurr and Oruue cotton Barru and SrNcn (1915) (1e75) citrus SeoeN.l and Keun (1974) taro Nemsup (1976) grapevines Senere and Salrosu tea KerHorsu (1929) (re77) Temo.r, (1922) Terooe et al. (1978) Israel citrus SHurov (1938) mulberry KevesHru.l (1 9 67, 19 7 2) apple M.qllsoun et al. cltrus Keruorsu (1979) orchards (1980a, b, c) NoHlnl and Yesuuet:su MeNsoun et al. (1981) (1965) orchards Hurusrrlre (1961) Japan rice Hauauum (1969,1971) Hurusrr're and Korcoo HesHruoro (1974) (te62) HrmNo and KrnrleNr Oruu.r, (1923) (1e76) Korea rice CHor et al. (1978) Irö er (1962) et al. (1976) Kxrve.^1. and Krnrtarr Horvo et al. (1978) (1e76) Oruue and Kru (1973) Karc and Krnrterr Pa.rr al. (1974) (1e78) Palr et mulberry Perx et al. (1973) Kavaxem (1975) Kavmem and Klnrlenr People's rice Anowvrrrous (1979) (1e75) Republic KavrHem er al. (1969, of China 1971) Krnrtexr (1977,1979) Philip- rice Gavenu and Ranos Krnr,reNr and K.crrve prnes (1e75) (1e7s) Innr Ann. Report (1923, Krnrrenr and Kavlsene 1974) (1e73) Innr Ann. Report (1926, Krnr:reNr et al. (1972) 1977) Koneve.snr (1975, 1977) Innr Ann. Report (1978) Korevasnr and Snrsara Lucrno and Rrnos (1e73) (re7s) Koveue (1972,1975, Serrles and Rrnos 1976) (1e75) Mrve: et al. (1978) Taiwan rice Cnru et al. (1974) Oxumr (1974,1977) Cnu and Oxuua (1920) Srsena and KrnrteNr Cnu and \(enc (1972, (1e74) 1973) Sesesl et aI. (1970) Cnu et al. (1975, 1976a, Sesese et aI. (1'973a,6) b' .) Suzurr and Krnrterr Cnu et al. (1977) (1e74) Tmnra (1973, 1975) Thailand rlce Orum (1968) Taraxa and Heua.uuro Oruue and'\$üoNcsrnr (1e68) (re73) available under natural conditions (MIvesHrre '1.968a, b; Hecsrnum 1970; Knssrrn 1973; BnnvrvrEyER and J62vrc 1975). Moreover' the prey composition established by means of prey preference tests often does not correspond to the prey composition realized in the field. Pest insects which are.regularly pr"y"d bn bv ä spider species in the laboratory may be entirely missing in the irrey co*positioä of the field (temporal andior sPatial isolation of-predator änd prey). Thus, the results of faunislic field studies and of nutritional analyses in the fabot^toty cannot be combined to a genuine whole. There is a great Spiders in natural pest control 325

Table 4. List of literature on the spider faunas in agroecosystems in Africa and in the Oceanic- Australian region

Country Agroecosystem Authors

Australia cotton Brsnor, (1979, 1980, 1981) Brsnop and Brooo (1980) Roou (1979) orchards Donnarn (1966) MeclrnaN (1923) Egypt clover Nncl" et al. (1975) cotton WrnsneNrv (1955) granati trees Tnurner (1981) Fiji Islands coconur Tolsnr et al. (1930) New Guinea coffee RosrNsoN and RorrNsox (1974) South Africa strawberry DrppnNe.tn-Scnorrvra.N (1976, 1979a, b) danger that speculations disregarding this ambiguity lead ro wrong results. Correspondingly, the opinions on the ecologicJl relevance of spidörs differ widely. Bnrstovr (1958) f.or instance estimates for Great Britain:-"Even if we were so cautious as to attribute a hundred victims in the year ro each spider, I calculate with a very conservative estimate of the spider popularion Tor the country as a whole that the weight of insects consumed innually in Britain exceeds that of the human inhabitants." The Canadian ecologist TunNsun (1973) give-s an even higher estimate: "I have calculated from 37 published censuses of spider populations that the mean density of spiders in a large variety of environments was 130.8 spiders/square meter, or l-308 OOO/hectare. If the food consumption of Argiope argentata were represenrarive of that of all spiders (which it is not), the average toial weight of foöd consumed per year by sbiders'would be ...''42490 kgThectare." f;.eferring to these coinpritation's Fonrrx (1982) wrote: "Even if such claims were realistic, it would be wrong to assume that spiders play a major role in the control of insect pests." In the following a survey is given about what is currendy-known on rhe ecological role of the spiders as predators of insects in undisiurbed grassland ecosystems, agroecosystems, and forest ecosystems (comp. also Vrrf 1953; KrncnNrn 1964, 1967; Luczer 1929; NvrrnLER and BrNz 1981a, 1982a; Nvpnrunn I982a, 1952b; RrrcHrnr 1984).

2 Ecological importance of the spiders in grassland and cereal ecosystems

Spid.ers-can- be of ecological importance in diverse respecrs. For example, they can be food for other predatorJ (\[Hrrcorrn 1974). Or the dead insects storea in spider webs can be food sources for kleptoparasites such as scorpionflies (TH.onnHrrr 1975; NvrnrrBn and BnNz 198öb). In this paper, only the role of spiders as predators will be discussed. An ecological importance of spiders as natural control agenrs is to be expected_in abandoned grasslands foremosr, where they can feid and repro- duce in the vegetation w:ithout human interference. Laige funnel web spiders and orb-weaving spiders often live in high populatiön densities in such habitats. Vith their strong webs they caprure large numbers of voluminous 326 M. Nyffeler and G. Benz insects. The orb-weaver Argiope bruennicbi, e.g. may caPture up to 7 grass- hoppers/web/day (NvrrnLER and BBNz 1981b). LoHI'trvnn and PnnrscHn'n Oiigl found on'the average 6 webs of A. bruenniclti per m2 in an abandoned grassland near Bonn (Fedäral Republic of Germany). This corresponds to a i'eb area of 3OOO m2lha ground aiea (Nxrnpr-rn 1982b). lt is evident that such a gigantic web area e"eris an enormous predatory pressure on insect pgPu-la- tioni. tf we consider that every spidef captures about 10 insects/web/day (Nvrreran 1982b), these orb-weaue.s *ould destroy 3 x 107 insects/ha in 50days. Polish studies have also shown that the orb-weaving spiders can exert a st.o.tg predatory pressure on the insect populations in the vegetation zone of abandonäd grassiands (Ifu;er et al. 1.968; Ke.ylx and OrncHovrcz 1970; Keler< 1971). Thus, web-building spiders eliminated 2540 % of the adult Diptera in undisturbed grasslands (Ke:er and Olr,cHovtcz 1970). Funnel web spiders (Gett,ts Agelena) can also occur in high population densities undiiturbed grasslands. Near Zurich (Switzerland), more than in -on 3 funnel web spiders /m2 the average were counted in some abandoned grassland habitäts. At certain points of aggregation within such a habitat, up to 12 funnel web spiders/m2 were found. The food of such funnel web spiders was composed of up to 23 "/" of bees (Nvnnnrnn and BrNz 1978)' In other- countries, too, it hafbeen observed that funnel web spiders can be predators of bees. Ornnnc (1960) wrote on this subject: "sometimes, 20 t_o 30 captured honey-bees on ihe average can be found in each web; thus, with a spider that frequent, a considerable-damage can result. This is why flocks of sheep are driven through the dangered aröas. The fleecy animals destroy the webs, which cannot be reilaced imäediately, as is the cäse in orb weaving-spiders." This example deÄonstrates that spiders can have a noticeable effect on insect -populations. Not only web-building spiders, but also hunting spiders (these are spiders which capture their prey without a web, e.g. wolf spiders) can be_important natural cöntrol age.ts of innertebrates in und-isturbed grasslands as the *ork of veN Hoox (1,971,) and ScHenrux (1971indicates. In contrast to undisturbed habitats, cultivated meadows and annual crops are in Central Europe unsuitable for the colonization by and survival of foliage-dwelling spidärs. In such agroecosysteTt +.. periodical.harvest of the vegetation is an extreme stress for ihe foliige-dwelling spiders (destruction of thä microhabitats and egg-sacs of numeroui spiders); a periodical recoloniza- tion of the fields from sirirounding undisturböd habitats is necessary (see also TrscHrnn 1958; Garrrn 1963; Lu"czer 1979). Population densitiei and bio- masses of spiders in the vegetation zone of cultivated meadows and cereal fields are low. As a consequence of the low spider densities, the numbers of orev killed bv the foliase'-dwelline spider communities of these cultivated i-t"b'it"tr lo'*, .o-p"rJd to the väuäs estimated for undisturbed grasslands; ir was estimared"r. that 2 2kginsects (fresh weigh)/ha/year are killed by spiders in the vegetation of such cultivated meadows and annual crops (see Ke;er 1971; Nvrrrlr,n and BnNz 1979a). The agroecosystems in parts of Southern Furope are to, some extent cultivatedin a considerably more extensive way than in Central Europe' It is therefore conceivable that ihe spiders still have ä gt"^tt predatory impbrtance in some cultivated fields of Soulhern Europe than in those of Central Furope. Thus, near Florence (Italy), rather high ipider numbers were found in the vegetation zone of a weedy cereal field (Srnnuu, pers. comm.). Spid.ers in natural pest control 327

In contrast to the vegeration zone, the spiders living on the ground surface of cultivated meadows and cereal fields in Central Europe show-relativelv high population densities (r-strategists , e.g. Erigorc otri1. Together wiih tf,e carnivorous beetles (Carabidae and Staphylinidae), they belöng to rhe most abundant ground-dwelling predatorr oT insects in ceräal fieldl near Zurich (Switzerland). German studies had also established that spiders form an important component within the ground-dwelling predator-cämplex (TrscH- rrn 1958, 1,965; Grnnn 1963; Besroov 1973; Bnassn 1,975; Besnoov and Mrnrr

3 Ecological importance of spiders in forest ecosystems and orchards

As a comparison to the studied Gramineae habitats, some tree ecosystems will be briefly discussed. \flith regard to forest ecosystems, VrronueNN (1978) attributes an important role to the ground-dwelling spiders as predators of insects, in combination with the other litter-dwelling predators. American scientists, too, concluded that the ground-dwelling spiders can be of great importance as predators of insects in forests (Crenkr and GneNr 1967; Mourorn and Rnrcnrs 1972: MeNny et al. 1976). In contrast to this, the predatory importance of the foliage-dwelling spiders of forest ecosystems is still controversial (Vn6 1953; KrncHnrn 1964;-NvFFELER 1982a). Some field studies had shown that certain spider species in the vegerarion were little effective in reducing populations of lepidopteran pesti (PorNrrNc 1966; KtRcunen 1967; Funure 1977). According to data by the last mentioned authors, some spider species could reduce moth populations by 5 "/" at the most. A slightly higher mortality was recorded by Encnr (1942), who 328 M. Nyffeler and G, Benz

estimated that during a pine moth calamity, 12-23 % of the moth population was destroyed by spiders; but at the same time he also observed that signifi- candy fewer moths were killed by spiders than by bugs. The ineffectiveness of certain spider species as predators of moths might be connected with the moths' ability to escape from these spiders'webs (KrncHNnn 1967; Nvrrnrnn and BnNz 1981c). However, this negative effect may not be very important, since other insects seem not to be killed at a higher rate by forest spiders. Thus, for a comparison, the mortality of chestnut gall wasp populations as a result of o/o) predation by spiders was of the same magnitude in normal years (7-2Q as that recorded by ENcnr for the pine moth (Nerallune and Nereuuxr 1977). Contrary to what has been said so far, some other authors have assumed that foliage-dwelling spiders may be important predators of Lepidoptera, aphids, and mosquitos in forest ecosystems (Sunr

4 Using spiders in biological pest control

Ground-dwelling spiders, together with the ground-dwelling raptorial Carabidae and Staphylinidae, form an important predator potendal in cereal fields. This predator potendal can be utilized within "programs for integrated pest control" (Besrnov and Mrrlre 1,977; KnnaNr 1979). Thus, attempts have been made in Japan to raise the spider density in rice fields artificially 6y releasing Drosopbila flies. This additional food then caused an increased fertility in the spiders (KorarasHt 1975). According to a report of the Chinese News Agency Xinhua of August 1,5,1979, in the Peoples Republic of China several species of spiders are introduced into rice fields as biological control agents of nce pests, As most spiders are cannibals, it would probably be difficult to breed large numbers of spiders in the laboratory. RunruRrsHoFEN (1976) and KRy,c.sHrMR (1967) have looked for an alternative. They have proposed that egg sacs of spiders could be gathered somewhere and placed in forests or mulberry plantations respectively to raise the population density of spiders. Another example on the use of spiders for biological pest control is reported from South Africa. There, spiders were settled in houses. SrnvN (1959) recorded a reduction of the fly populations by 99 % within 2r/z months and, at the same time, a pronounced decrease of gastrointestinal infections of men in that region, because the vectors of disease were destroyed. There are several reports in literature about house-dwelling spiders as predators of pest insects (comp. Lonauoo 1,929; Matnrs and BrnreNo 1933; Hasn 1934; Saurnr 1936; Kunrr.raNN 1970/

I Spiders in natural pest control 329

71 ; Eowanns 1979); in stables in Canada it could be observed that th_e prey of the comb-footed spider Steatoda biponaoto consisted mainly of harmful insects such as house ilies, Tenebrio beetles, etc. (Nvrrrrrn unpubl.). 'What about the possibility of utilizing spiders for biological pest control in Central Europe? Hire, spidärs could only-become of greater impo-rtance, if it were oossiblä to increase the numbers of spiders in cultivated fields. This *ould be possible if the area of the "ecologiial cells" (abandoned grasslands, hedges, *ät areas, etc.; definition of "ecological cell" see Klorr 1978) which servi as predator-reservoirs for agroecosystams, could be enlarged. However, uncultivited lands of that kind may serve as reservoirs for predators and pest organisms as well. The expansion-of the area of the."ecolögical cells" could thärefore also bring aboui an increase of pest incidence. \Vith present day- knowledge it is difficult to estimate, whether the useful effect (reservoirs of predatori) or the detrimental effect (reservoirs of pests) would ultimately such "ecological prevail. Intensive research on the complex functions of -ce-lls" in the agricultural landscape would be-necessary, before the expansion of their area wit-h the intentions oT increasing the predators' densities in the culdvated areas could be recommended.

Acknowledgements This research was made possible by a predoctoral fellowship of the Swiss National Science Foundation (grant 3.0020'.76). Ve ihank Prof. Dr. SürNrInio L. Srrnr-rrc, Department of Entomology, Texas A&M University (College Station/USA), for his review of this paper.

Zusammenfassung

Spinnen in der natürlichen und biologischen Schädlingsbekämpfung: Ein Literaturüberblich Veltweit wurden in den letzten 20 Iahren in verschiedenen Ökosystem-Typen zahlreiche Studien über die ökologische Bedeutung der Spinnen als Prädatoren durchgeführt. Trotz der großen Zahl durchgeführteispinnenökologiicher Studien ist die Funktion der Spinnen als Prädatoren von Insektän und anäeren lnvertäbraten heute noch weitgehend ungeklärt. ln vorliegender Arbeit wird die Literatur über dieses Thema (insgesamt sind 3Ob wissenscfiaftliche Arbeiten. die zwischen 1920 und 1984 über dieses Thema publiziert wurden, aufgeführt) analysiert. Mehrere europäische und amerifranische.studien haben gezeigt,daß Spinnen_in unbewirtschaf- teten Grasland-Ökoryrte-e.t und Forst-Ökosystemen eine gioße ökölogische Bedeutung als Prädatoren von Insekien haben können. Auch in ungespritzten Obstanlagen, die man bis zu einem gewissen Grad mit Forst-Ökosystemen vergleicherikann, stellen Spinnei manchmal eine häufige Frädatorengruppe dar. Im Gegensatz dazu-ist die ökologische Bedeutung der in Kulturfeldern Iebenden Söin;än als lnsekteniertilger noch weitgehend ümstritten. Einige europäische Studien deuten daräuf hin, daß den in der Vegetationsschicht von Kulturfeldern lebenden Spinnen ihrer niedrigen Populationsdichten wegen wahrscheinlich keine große Bedeutung als Prädatoren von lnsektin zukämmt. Aus anderen-europäischen Studien gehihervor, daß die epigäischen Spinnen der Kulturfelder bezüglich ihrer Abundanz eine dominänte Prädatorengruppe darstellen, deren Funktion als Kleinräuber allerdings noch weitgehend unbekannt ist. In asiatischen Reisfeldern, die nicht oder wenig mit Pestiziden Sehandelt wurden, können Spinnen eine wichtige Bedeutung als Prädatoren von-Schädlingen haben. Reisfelder sind Sumpfökolysteme; auch in europäischen_und amerikanischen Sumpfökäsystemen können Spinnen eine abundante Prädatorengruppe dqrstellen. In Südafrika *rriden Spinnen erfolgreich- bei der biologischen Bekämpfung von Fliegen in Häusern eingesetzt.

References

Auoxvuous, 1979:Trials on the protection and use of the spiders in large paddy fields. Biology Dept., Hunan Teachers' Colt., Changsha, Hunan, P. R. China (in .Chinese)' IRR Inititute, 1973-1.978: Ann. Report {or 't972-1977. Los Bafros, Philippines. Acurran, P. G., 1965: Notas sobre-las aranas en el campo cultivado. Rev. Peruana Ent. 8, 81-83. 330 M. Nyffeler and. G. Benz

1974: Aranas del campo cultivado l: poblacion de araneidos en algodonales de Canete, - - Huaura y Rimac. (Spiders of crop fields - l: population of araneids in corton plantations in Canete, Huaura and Rimac). Rev. Peruana Eit. 17,21,-27. Arvqutsr, S., 1981: Spiders and harvestmen in strawberry fields in sourhern Sweden. Ent. Tidskr. 102, 159-162. Asnrxrettv, N. 2., 1973: The life forms of spiders (Araneae) inhabiting wheat fields in the Kustanay region. Entomol. Rev. J2, 335-341. Batrv, C. L.; Cnaoa, H. L., 1968: Spider populations in grain sorghums. Ann. Ent. Soc. Am. 61, 567-571,. Barocrr, J.; Lorsa, J., 1956: Untersuchungen über die Zoozönose des Luzernefeldes. Acta zool. acad. sci. Hung. 2, 17-714. Bannosa, J. T.; Rrlceoo, G. M.; Frrrro, M. L.,7979: Population fluctuation of sugarcane froghopper, Mabanaroaposticatastal,lS55 and its natural enemies in Campos, R.J.,in1977. An. Soc. Entomol. Bras. 8,3946. Besroow, T., 1973: Der Einfluß epigäischer Raubarthropoden auf die Abundanz phytophager Insekten in der Agrarlandschaft. Pedobiologia, Jena, 13, 41,V22. Basnoov, T.; Mrrrrn, H., 1977: Aspekte der Parathion-Anwendung in Weizenfeldern. Nach- richtenbl. Deut. Pflanzenschutzd. 29, 65-69. Berru, G. S.; SINcH, 8.,7975: A note on spiders predatory on the insect pests of cotton. Sci. and Cub. 41,212-21.4. BrNz, G.; Nvrrnun, M., 1980: Ecology of spiders in meadows nea;' Z,trich (Switzerland). Proc. Int. Arachnol. Congr., 8th, Vienna, L21,-1,25. Bnvnn, R., 1981.: Zv Dynamik der Spinnen- und Weberknechtfauna auf einer Kulturfläche mit wechselndem Pflanzänbestand im Verlaufe von 5 Jahren im Raum Leipzig. Faun. Abh. Mus. Tierk. Dresden 8, 1,1,9-1,30. BrrsrNc, S. V., 1920: Quantitative studies in the food of spiders. Ohio J. Sci. 20,215-260. Brsnoe, A.L.,1979: Theroleof spidersaspredatorsinacottonecosystem.M.Sc.thesis,Univ. Queensland. 1980: The composition and abundance of the spider fauna in south-east Queensland cotton. - Aust. J. Zool. 28,699-708. l98l: The spatial dispersion of spiders in a cotton ecosystem. Aust. J. Zool. 29, 15-24. -Btsxon, A. L.; Br-ooo, P. R. B., 1980: Arthropod ground strata composition of the cotton ecosystem in south-eastern Queensland, and the effect of some control strategies. Aust. J. zool. 2g, 693-697. BLlcrntrsterr, C. C. ; Huccers, J. L., 1962: Soybean insects and related arthropods in Missouri. Univ. Missouri Agric. Exp. Srn. Res. Bull. 803. (Cit. SüHrtrrn, A. G., 1973.) Boucss, M.,.. 1953: Die Fauna der 'Wiesen unter besonderer Berücksichtigung der Mahd. Z. Morph. Ökol. Ti... 42,225-277. 1958: Biorönotische Untersuchungen- über die Tierwelt von Klee- und Luzernefeldern. Z. - Morph. Ökol. Tiere 47,309-373. BosrreNs, R.; Corrrnrr, M. P., 1977: Araign6es rares ou nouvelles pour la faune belge des cultures. Bull. Ann. Soc. r. belge Ent. 113, L00. Bnessn, D., 1,975: Die Arthropodenfauna von Getreidefeldern auf verschiedenen Böden im Braunschweiger Raum. Pedobiologia, lena, 15, 40541,4. Bnryunnn, A., 1.967: Preliminary data for estimating the biological production of wandering spiders. In: Secondary productivity of terrestrial ecosystems. Ed. by K. Prrnusrwrcz. pp. 821-834. \Warszawa, Krakow. 1978: Analysis of the trophic stmcture of some grassland ecosystems. Pol. Ecol. Stud. 4, - 55-128. Bnrrrr,rtvnn, A.; JdZvrr, J., 1.975 Consumption of wandering spiders (Lycosidae, Araneae) esdmated in laboratory conditions. Bull. Acad. Pol. Sci., Cl.lI,23,9J-99. Bnrsrovn, V. S., 1958: The Vorld of Spiders. London: Collins. Bunrercn, J. G.; Youxc, J. H.; MonnrsoN, R. D., 1973: Strip-cropping's effect on beneficial insects and spiders associated with cotton in Oklahoma. Environ. Entomol. 2,281-285. Cennor.r, D. P., 1980: Biological notes on the spiders of some citrus groves in central and southern California. Ent. News 91,1,47-1,54. Cuarr, D. A., 1956: Predaceous spiders in orchards in south-eastern England. J. Hort. Sci. J1, 35-_4,6. CnauvrN, R., 1960: La faune du champs cultiv6 et surtout du champ de luzerne. Revue Zool. agric. 59, 1,4-37. 7967 Die Velt der Insekten. München: Kindler. -CHrnnnrr, M,1,964: The distribution of spiders on the Moor House National Nature Reserve, 'Westmorland.J. J. Anim. Ecol. 33, 2748.

- Spiders in nataral pest control 331

Cnru, S. C.; Crru, Y. I.; Lurc, Y.H., 1974: The life history and some bionomic notes on a spider, Oedothorax insecticeps Boes. et St. (Micryphantidae: Araneae). Plant Prot. Bull., Taiwan, 16, 1.53-161.. Cnor, S. Y.; Lnr, H. R.; Rvu, J. K., 1978: Effects of carbofuran root-zone placement on the spider populations in the paddy fields. Korean J. Plant Prot. 17,99-1.03. Cnu, Y.; OrÜue, C., 1970: Preliminary survey on the spider fauna of the paddy fields in Taiwan. Mushi, Fukuoka, 44, 65-88. Crru, Y. I.; WeNc, C.T.,7972: Studies on the Llcosa pseudoannulata (Boes. et Str.) (Araneae: Lycosidae). (I) External features and habits. Plant Prot. Bull., Taiwan, 14,169-174. CHu, Y. I.; Sfexc, 5.T., 1973: Studies on the feeding habits of Lycosa pseudoannulata (Boes. et Str.). Plant Prot. Bull., Taiwan, 15, 13-20. Cnu, Y. I.; Ho, C. C.; CnrN, 8.J., 1975: Relative toxicity of some insecticides to green rice leafhopper, brown pianthopper and their predator Lycosa pseudoannulata. Plant Prot. Bull., Taiwan, 17,424-4'30. 7976a: The effect of BPMC and Unden on the predation of Lycosa spider (Lycosa pseudoannulata-). Plant Prot. Bull., Taiwan, 18,42-57. CHu, Y. I.; LrN, D. S.; Mu, T., 1976b: Relative toxicity of 9 insecticides against rice insect pests and their predators. Plant Prot. Bull., Taiwan, 18,369-376. L976c:The effect of Padan, Ofunack and Sumithion on the feeding amount of Lycosa pseud.oannulata (Boes. et Str.) and Oedothorax inseaiceps Boes. et Str. (Lycosidae and Micryphantidae, Arachnida). Plant Prot. Bull., Taiwan, 18,377-390.

with the effect of Bidrin on rhe extent of feeding by Lycosa pseudoannulata and Oedothorax inseaiceps. Plant Prot. Bull., Taiwan, 19, 1-12. Crenr, E. \(/.; Gucr, P. A., 1961: Some predators and scavengers feeding upon pink bollworm moths. J. Econ. Ent. 54, 81,5-816. Crenrr, R. D.; GneNr, P. R., 1967: An experimental study of the role of spiders as predators in a forest litter community. Part l. Ecology 49, 1152-1.154. CortrNrr, P.; or Crrncq, R., 1977: Study of the arachnofauna in fields of winter wheat. Parasitica 33, 1.38-147. CuuN, J. D.; Rusr, R. V., 1980: Comparison of the ground surface and foliage dwelling spider communities in a soybean habitat. Environ. Entomol. 9, 577-582. Czeyx,l., M.; Goos, M.,7976: The spiders (Aranei) of sugar-beet fields in Pawlowice'Wielkie near '\(i roclaw. Pol. Pismo Ent. 46, 1,79-'1,85. Czeyxe., M.; K-e.rre, C., 1976: Spiders (Aranei) in potato agrocoenosis in Pawlowice Süielkie near '\(i roclaw in 1,971,-1,974. Pol. Pismo Ent. 46, 623-629. DasnovsKA-Pnor, E.; LuczAK, J.; TenvIo, K., 1968a: The role of prey and predator density in the process of mosquito reduction by spiders in field experiments. Ekol. Pol. A 16, 1-81. 1968b: The predation of spiders on forest mosquitoes in field experiments. J. Med. Ent. 5,252-256. DnaN, D. A.; Srrnurc, W. L.; HonNrR, N. V., 1982: Spiders in eastern Texas cotton fields. J. Arachnol. 10,251-260. nr Crrncq, R., 1979: On the influence of the soil fauna on the aphid population in winter wheat. Int. Symp. "Integrated Control in Agriculture and Forestry", Vienna, 8-12 Oct. Summaries of main lectures and discussion papers. p. 79. Dr:rz, L. L.; vel DuvN, J. V.; Bneorrv, J. R.; Renn, R. L.; Bnoors, \(. M.; SrInNan, R. E., L976: A guide to the identification and biology of soybean arthropods in North Carolina. Techn. Bull., North Carolina, Agric. Exp. Stn. No. 238. Drrcnrv, K.; Keler, A., 1974: Analysis of a sheep pasture ecosystem in the Pieniny Mountains (the Carpathians). XVI. Effect of pasture management on rhe number and biomass of spiders (Araneae) in two climatic regions (the Pieniny and the Sredna Gora Mountains). Ekol. Pol. 22, 693-71,0. Drnlnrcn, 10(.; Görzs, R.,7974: Ökologische Untersuchungen der Spinnenfauna einer Agro- biozönose (unter besonderer Berücksichtigung der Araneae). Unpubl. Dipl.-Arb., Sektion Biowissenschaften, Fachbereich Zoologie, Martin-Luther-Univ., Halle. DtNcrtn, M., 1935: Die Tierwelt des Spargelfeld es. Z. ang. Enr. 21 , 291-327 . DrppnNeen-ScHoEMAN, A. 5., 1976: An eCological study of a spider population in strawberries with special reference to the role of Pardosa crassipalpis Purcell (Araneae: Lycosidae) in the control of Tetranychus cinnabarinus (Boisduval). M. Sc. thesis, Rand Afrikaans Univ. 1979a: Spider communities in strawberry beds: seasonal changes in numbers and species - composition. Phytophylactica 1 1, 14. 1979b A simple te-hnique for studying feeding behaviour of spiders on mites. Bull. Brit. - Arachnol. 9oc.4,349. 332 M. Nyffeler and G. Benz

DoaNr, J. F.; Donoerr, C. D., 1979: Seasonal captures of spiders (Araneae) in a wheat field and its grassy borders in central Saskatchewan. Can. Ent. 111, 439445. DoNoer-n, C. D., 1956: Annotated list of spiders (Araneae) from apple trees in Nova Scotia. Can. Ent.88, 697-700. 1958: Note on population densities of spiders (Araneae) in Nova Scotia apple orchards. Can. - Ent.90, 111-113. 1966: The spider fauna (Araneida) of deciduous orchards in the Australian capital territory. - Austr. J. Zool. 14,1157-1192. 1971: Spiders of Heasman's field, a mown meadow near Belleville, Ontario. Proc. Ent. Soc. - Ontario 101,62-..69. 7977: Life histories and distribution patterns of hunting spiders (Araneida) in an Ontario - meadow. J. Arachnol. 4,73-93. DoNoarr, C. D.; BrnNs, M. R, 1977: Effect of weather factors on spiders (Araneida) in an Ontario meadow. Can. J. Zool. t5, 1336-1.341.. DoNoere, C. D.; Rrnwrn, J.H.; Sruprr, R.8., 1972: Diel activity periodicities in meadow arthropods. Can. J. Zool. t0, 1155-116J. Doroeu, C. D. ; Penrrr, B.; Prrnn, D., 1979: A 6-yeer study of spiders (Araneae) in a Quebec apple orchard. Can. Ent. 111,377-380. Doroerr, C. D.; RroNrn, H.; Fannerl, E.; Snlrlnrr, R. B.; TunNnuu-, A. L., 1970: 'lü(/andering J. of hunting spiders in a meadow. Bull. Mus. Hist. nat., Paris, 2" s6r., 41 (suppl. 1), 6144. Donnrs, P. R., 1970: Observations on the impact of certain insecticides on spider populations in a cotton field. Proc. Ark. Acad. Sci. 24,53-54. Durrnv, E., 7962: A population study of spiders in limestone grassland. J. Anim. Ecol. -?1, 571,-599. 1974: Comparative sampling methods for grassland spiders. Bull. Brit. Arachnol. Soc. J, - 34-37. DuNN, J. A., 1,949: The parasites and predators of potato aphids. Bull. Ent. Res. 40,97-1.22. Eowenis, C. A.; Burrrn, C. G.; Lorrv, J. R., 1976t The invertebrate fauna of the park grass plots. II. Surface fauna. Rep. Rothamst. exp. Stn. 1975, Part2, 63-89. Eowanns, G. 8., 1979: The giant crab spider, Heteropoda oenatoria. Florida Dept. Agric. Entomol. Circ. No. 205. Ercrr, H., 1942: über die Populationsbewegung des Kiefernspanners (Bwpalus piniariws L.) in verschiedenen Bestandestypen. Z. ang. Ent. 29, 116-163. Evnnrv, R. T., 1938: Spiders äird insects Iound associated with sweet corn with notes on the food habits of some species. I. Arachnida and Coleoptera. Ohio J. Sci. J8, 134-140. Fonrrx, R. F., 1932J Biology of Spiders. Cambridge, Mass., and London; Harvard Univ. Press. Fox, C. J. S.; DoNoerr, C. D., tgTZ: Annotated list of spiders (Araneae) from hayfields and their margins in Nova Scotia. Can. Ent. 104, 1911-1915.

Entomol. Soc. -1-1, 241.--243. Fnesnn, A. M., 1982: The role of spiders in checking cereal aphid numbers. Ph.D. thesis, Univ. East Anglia (in prep.). Funure, K., 1977: Evaluation of spiders, Oxyopes sertatas md O. badius (Oxyopidae) as a mortality factor of gypsy moth, Lymantria dispar (Leptdoptera: Lymantriidae) and pine morh, Dendrolimus Eectabilts (Lepidoptera: Lasiocampidae). Appl. Ent. Zool. t2, 313-324. Gerrcxa, 8.,1966: The role of predators in the reduction of two species of potato aphids' Aphis nasturtä Kalt. and A. frangulae Kalt. Ekol. PoL. A 14,245174. Gancre, R. ; ScsrrNcrn , E. 1., 7972: Studies of spider predatio n on Aedes dorsalis (Meigen) in a salt marsh. Proc. and Papers of 40th Ann. Conf. Calif. Mosquito Control Ass., Jan. 3l-Febr. 1-2, 1972. pp.117-118. Gavanna, M. R.; Ranos, R. S., 1975: Studies on the biology of the predatory wolf spider Lycosa

pseudoannulata Boes. et Str. (Araneae: Lycosidae). Philipp. Ent. 2, 427444. , Gelr-En, H., 1963: Die Spinnen- und Weberknechtfauna nordwestsächsischer Felder (Die Everte- bratenfauna mitteldeutscher Feldkulturen Y). Z. ang. Zool. 50,257-272. Goos, M., 1973: Influence of aphicides used in sugar-beet plantations on arthropods. II. Studies on arachnids - Arachnoidea. Pol. Pismo Ent.43,851-859. Hacsrnuu, D. \ü., 1970: Ecological energetics of the spider Tarentula kocbi (Araneae: Lycosidae). Ann. Ent. Soc. Am. 63,1297-1304. Heuar,runa, T., 1969 Seasonal fluctuation of spider population in paddy fields. Acta Arachnol. 22, 4o-50. 1971: Ecological studies of Pirata subpiratlcas (Boes. and Strand) (Araneae, Lycosidae). - I. Acta Arachnol. 23, 1,1-1,8. Spiders in natural pest conüol 333

HennrsoN, J. O., 1968: Some spiders associated with banana plants in Panama. Ann. Ent. Soc' Am. 51,878-884. Hesr, A., 1934: Über eine erfolgreiche biologische 'Wanzenbekämpfung durch die Spinne Tbanatus flavidzs. Naturwissenschaften 22, 649450. Hesrrruoro,'R., 1974: Some spiders of the paddy field and their seasonal fluctuation. Agric. Chem. 21,22-26. HrNsrrr, S. D.; Lorc, !ü. H.; Ronov, L. R.; McConr'rrcr, V. J.; CoNcrrNNr., E. J., 1'961: Effects of insecticides on the predaceous arthropod fauna of Louisiana sugarcane fields. J. Econ. Ent. t4,146-149. HrvonuenN, B., 1953: Agrarökologische Problematik, dargetan an (Jnterslchungen über die Tierwelt der Oberflächä der Kulturfelder. Unveröffentl. Diss., Univ. Kiel. (Cit. Gnrun, H., 1e63.) Hrnexo, C.; Krnrrexr, K., 1976: Paddy ecosystem affected by nitrogenous fertilizer and insecticides. In: Science for better environment. Proc. of the Intern. Congr. on the Human Environment, Kyoto, 1,975. Tokyo: Asahi Evening News. Horvo, N.; Las, M. u.; Panr, j. 5., 1976: Some aspects of population dynamics of rice leafhoppers in Korea. Korean J. Piant Prot. 15, 1.1.1-126. HonNnn, N. Y.,1972: Metaphidippus galathea as a possible biological control agent. J. Kans. Ent. 9oc.45,324-327. Hovann, F. rtrü.; Or-rven, A. D., 1978: Arthropod populations in permanent pastures treated and untreated for red imported fire ant control. Environ. Entomol. 6,901-901. Hovnrr-, J. O.; Prrrrowsrl, R. L., 1971 : Spider populations in alfalfa, with notes on spider prey and effect of harvest. J. Econ. Ent. 64,163-168. HuHre, V.; RaerratNrN, M., 1974: Spider communities of leys and winter cereal fields in Finland. Ann. Zool. Fenn. 11,97-1.04. Hurusrue, S., 1961 : Studies on the insect association in crop field. XXI. Notes on spiders in apple orchards. Jap. J. appl. Ent. Zool. 5,270-272. HurÜsrua, S.; Kor.lob, f., lgtz: Further evaluation in the feeding potential of the predaceous insects and spiders in association with aphids harmful to apple and pear growing and the effect of pesticides on predators. Jap. J. appl. Ent. Zool. 6, 274-280. lrö, J.; MlrasHlra,-K.; Srxlcucnr, K- 1962: Studies on the predators of rice crop insect pests, using the insecticidal check method. Jap. J. Ecol. 12, l-1.1.. JonNsoN, E. K.; Yourc, J. H.; Morren, D. R.; MonnIsoN, R., 1976: Effects of three insect control schemes on populations of cotton insects and spiders, fruit damage, and yield of Vestburn 7O cotton. Environ. Entomol. ,, 508-510. Jurnnr, J. A., 1961: Observations on arthropod predators of the European pine shoot moth, Rbyacionia buoli.ana (Schiff.) (Lepidoptära: Olethreutidae), in Ontario. Can. Ent. 9J, 195-1.98. KecrN, M., 1943: The Araneidae found on cotton in Central Texas. Ann. Ent. Soc. Am. 36, 257-258. KarHorsu, K., 1979: Spiders and their seasonal fluctuation in a citrus orchard and a tea field in Mie Prefecture. Atypus 74,29-39. Ke,1ex, A., 1960: Chinges in the abundance of spiders in several meadovrs. Ekol. Pol. A I, 1,99-228. 7962: Comparison of spider fauna in artificial and natural meadows. Ekol. Pol. A 10, l-20. - 1971: Productivity invesdgadon of two types of meadows in the Vistula Valley. IX. Produc- - tion and consumption of field layer spiders. Ekol. Pol. A 19, 197-211. 1978: The effect öf fertilizers onnumber and biomass of spiders in a meadow. Symp. zool. - Soc. London 42, 125-129. 1980: Do the changes caused in spider communities by the application of fertilizers advance - with time? Proc. Int. Arachnol. Congr.,8th, Vienna, pp. 1l!-1 19. Keyer, A.; Jerurczvr, H., 1975: Experimental studies on ipider predation. Proc. Int. Arachnol. Congr., 6th, Amsterdam, 1974, pp.82-85. Kayer, A.; Orncnowrcz, 8., 1969,1910:The role of web spiders in elimination of Diptera in the meadow ecosystem. Bull. Mus. Hist. nat., Paris, 2" s6r., 4.1 (suppl. 1,),233-236. Kalar, A.; Aronzrlrvsxa, L.; WoJcIx, 2.,7968: The role of spideis in the decrease of damages caused by Acridoidea on meadows - experimental investigations. Ekol. Pol. A 16,755-764. K,t1ex, A.; Iinrrurrun, A.; Prrer, J., 1971 : Productivity inveltigation of two types of meadows in the Vistula Valley. Predatory arthropods. Ekol. Pol. A 19,223-233. Kerrvn, N.; Krnrrnxr, K., 1976: Compariion of sampling techniques to estimate the population density of Lycosa pseudoannulata in a paddy field. Jap. J. appl. Ent. Zool. 20,21-25. Karoor, M.8., 1976: Brown planthopper in rice and its control. Indian Farming 27,3-5. 334 M. Nyffeler and G. Benz

KeNc, J.; Krnr'rrNr, K.,7978: \(i inter mortality of the green rice leafhopper (Nephotettix cinaiceps Uhler) caused by predation. Jap. J. appl. Enl Zool. 20, 21.-25. Kawenanr, 5., 19i54 Population-dynamlcs of micryphantid spiders in the paddy field. Bull. Kochi Inst. Agr. & Forest Sci., No. 7,53-64. Ke.wanenr, S.; KrnrreNr, K., 7975: Survival and egg sac formation rates of adult females of Lycosa pseudoannulata (Boes. and Str.) (Araneae, Lycosidae) in the paddy field. Appl. Ent. Zool. 10,232--234. Kewenena, S.; KrnrraNr, K.; Sesane, T., 1971: The selective activity of rice-pest insecticides against the green rice leafhopper and spiders. Botyu-Kagaku 36,121-128. (Cit. KrnrreNI, K., 1977.) Ke.wanana, S.; Krnrreu, K.; Sasene, T.; Neresuyr, F.; Oruue, C'7969: Seasonal changes in abundance and the faunal composition of spiders in the paddy field, with special reference to their relation to the seasonal prevalence of the green rice leafhopper, Nepbotettix cincticeps Uhler. Proc. Assoc. Pl. Prot. Sikoku 4,3344. Kev,rsnrue, I., 1960: Studies on spiders as natural enemies of crop pests (I). Daily activities of spiders in the cabbage fields, esiablishment of spiders liberated in the fields and evaluation of the effectiveness of spiders against crop pests. The Science Bull., Fac. Agric. Kyushu Univ. 18, 1-24_ 1967: Study on spiders (particularly referring to grass-spiders) to prey upon fall-web worms - (Hyphantrya cunea Drury). Acta Arachnol. 21, 1-30. 7972: Study on grass spider as a predator to Hypbantria cunea Drury (Experiment on - effectiveness as a predator oI Agelena opulenta L. Koch). Acta Arachnol. 24,6V72. Krssrrn, A., 1973: A-comparative sludy of the production of eggs in eight Pardosa species in the field (Araneida, Lycosidae). Tijdschr. Entom. 116,2341. KrncnNnn, \(., 1964: Bisher Bekanntes über die forstliche Bedeutung der Spinnen - Versuch einer Literaturanalyse. \üaldhygiene S, 161-198. 1.967: In welihem Maße'bieinflussen Spinnen- die Mortalitätsrate des EichenwicUers (Tortrix - iiridono L.)? Valdhygiene 7,23-31. KrnrtaNr, K., 1977l. Systems approach for managernent of rice pests. In: J. S. Pecxnx.; \?Hrrr, D. (Eds.). Proc. XV, Int. Congr. Entomol., Washington, D.C., August 19-27,1976, Entomol. Soc. Am., Md., pp. 591-598. 1979: Pest management in rice. Ann. Rev. Ent. 24, 279-312. -Krnrlert, K.; K.*rre, N., 1975: An analysis of the predator-prey system in the paddy field. Res. Popul. Ecol. 17,29-38. KinrreNr, K. ; K.r.venene , 5., 1973: Food-chain toxicity of granular formulations of insecticides to a predator, Lltcosa pseadoannulata, oI Nepbotettix cincticeps. Botyu-Kagaku 38, 69-75. (Cit. KrnrraNr, K., 1.977.) Krnrrerr, K.; K.r,veH,Lne, S.; Sesare, T.; Neresulr, F., 1972: Quantitative evaluation of predation by spiders on the green rice leafhopper, Nepbotettix cincticeps Uhler, by a sight- count method. Res. Popul. Eco[. 1.], 187-200. Krnr.r, B., 1978: Untersuchung der epigäischen Spinnenfauna (Araneae) in Rebkulturen. Staats- examensarbeit in Zoologie, Univ. Freiburg i. Br. Kr-orr, \iü., 1978: Ökologie"der Tiere. Stuttgärt: Ulmer. Konevnsnr, S., 1975: The effect of Drosophila release on the spider population in a paddy field. Appl. Ent. ZooL 10,268-274. 1977: Change in population density of spiders in paddy field during winter. Acta Arachnol. 22 - (special no), 247:251. KonavnsHr, S.; SHrn,Lra, H., 1973: Seasonal changes in population density of spiders in paddy fields, with reference to the ecological control of the rice insect pests. Jap. J. appl. Ent.Zool. 17, 193-202. Kovar, G., 1976: Biologische Bekämpfung des Kartoffelkäfers. 4Zrscita Rastenij, Moskau, 21, 29. Kovena, J., 1972: Spiders in paddy fields in Akita Prefecture. Ann. Plant Prot. North Japan 23, 127. 1975: Studies on the diminution of insecticide application to the rice xem borer, Cbilo - sappressalis Valker. III. The effect of insecticide application on the density oflarvae of the rice stem borer and spiders. Jap. J. appl. Enr. ZooL 19, 125-132. 1976: The effects of low concentration of chlorphenamidine on the rice stem borer, Cbilo - suppressalis lValker, and on spiders. Rice Entomol. Nev/sl. 4, 29-30. Knerunn, P., 1,961,: Einfluß einiger Arthropoden auf Raubmilben (AcÄ). Z. ang. Zool. 48, 257-31.1. KurrueNr, E., 1,970/71: Bemerkenswerte Konvergenzen im Verhalten cribellater und ecribellater Spinnen. Freunde des Kölner Zoo 13,123-150. Spiders in natural pest control 335

LncNrn, E. F.; Oarrr,rax, E. R., 1964: Spiders on apple in S7isconsin and their abundance in a natural and two artificial environments. Can. Ent. 96,1202-1207. Lnrcrr, T. F.; Hurrpn, R. E., 1969: Predaceous spiders in California cotton. Calif. Agric.23,4-5. Lrsan, C. D.; Unzicker,J.D.,7978a: Soybean spiders: species composition, population densities and vertical distribution. Ill. Nat. Hist. Surv. Biological Notes 707. 1,978b: Life history, habits, and prey preferences of Tetragnatba laboriosa (Araneae: - -Tetragnathidae). Environ. Entomol. 7, 879-884. Locxar, G. H., 1978: The Leckford survey spiders progress reporr.Ja;n.1978. Locrrnv, T. C.; Srrarrn,J. \(.; Scorr, W. P.; Penrncre, C. R., 1979:Popriation fluctuations of two groups of spiders from selected cotton fields in Panola and Pontotoc Counties, Mis- sissippi, 1977. Southwest. Entomol. 4? 2U24. '\üildstauden- LoHrtanvan, lV.; Pnrrscnrn, P., 7979: Uber das Zustandekommen halbruderaler Quecken-Fluren auf Brachland in Bonn und ihre Bedeutung als Lebensraum für die \üespen- spinne. Natur und Landschaft 54,253-259. Lonalno, N. T., 1929: A biological method of destroying bedbugs. Sci. Monthly 29,265-268. Lucrno, R. M.; Renos, R. S., 1925: Toxicity of Carbofuran and Gamma-BHC to the spiderlings of the wolf spider, Lycosa pseudoannulata Boes. et Str. (Araneae: Lycosidae). Philipp. Ent.3, 69-75. Luczer, J., 1968: An attempt at quantitative estimation of some factors on the predator-prey system, on the example of spiders and mosquitoes. Ekol. Pol. B 14, 343-350. 1,974: Ecological groups of spiders of potato and rye fields. Bull. Acad. Pol. Sci., Cl. II, 56r. - Sci. biol. 22,377-383. 1,975: Spider communities of the crop-fields. Pol. Ecol. Stud. 1, 9J-1,1,0. - 1976: (A decrease of arachnofauna in agrocoenoses as a result of agrotechnical treatments). In: - Entomologia a ochrona srodowiska. Ed. by H. S.qNoNrn. Warszawa: P\7N. pp. 99-101. 7979: Spiders in agrocoenoses. Pol. Ecoi. Stud. J, 151-200. -LuneN, U., 1979: Tagesrhythmik und Jahresrhythmik epigäischer Arthropoden (insbesond. Aranei, Carabidae) einer mesophilen \üiese des lnnsbrucker Mittelgebirges (Rinn 900 m NN, Osterreich). Unveröff. LA-Hausarbeit in Zoologie, Univ. Innsbruck. Macl-rrr.lx, C. R., 1973: Natural enemies of the light brown apple moth, Epipblas postoittana, in the Australian capital territory. Can. Ent. 105, 681-700. Marrnr, G. V. ; Burcnrn, J. V.; Zrsrc, M., 7976: DDT transfer and metabolism in a forest litter macro-arthropod food chain. Pedobiologia, lena, 1 6, 81,-98. MaNsoun, F.; Rosrx, D.; Snurov, A., 1980a: Functional response of the spider Chiracantbiurn mild.ei (Arachnida: Clubionidae) to prey density. Entomophaga 25,313-316. 1980b: A survey of spider populations (Araneae) in sprayed and unsprayed apple orchards. Acta Oecologica, Oecol. Applic. /, 189-197. 1981: Disturbing effect of a spider on larval aggregations of Spodoptera littoralis. Enr. exp. & appl. 29,234-217. MeNsoun, F.; Rosrr,r, D.; Snurov, A.; Pmur, H. N., 1980c: Evaluation of spiders as biologicai control agents of Spodoptera littoralis larvae on apple in Israel. Acta Oecologica, Oecol. Applic. 7, 221232. Maturs, C.; Brnreno, L., 1933: Une araign6e africaine: Plexippus paykulli, enemie naturelle des Stegomyia hötes des maisons. C. R. Acad. Sci., Paris, 197,271-272. Maunrn, R., 1975: Die Vielfalt der Käfer- und Spinnenfauna des \üüiesenbodens im Einflußbereich von Verkehrsimmissionen. Oecologia, Berl., 1 4, 327-351. 1975: Epigäische Spinnen der Nordschweiz L Mitt. Schweiz. Ent. Ges. 48,357-376. -McCannnnv, J. P.; HonsruRGH, R. L.,7978: Laboratory feeding studies with selected spiders (Arachnida: Araneae) from Virginia apple orchards. J. New York Entomol. Soc. 86, 308. 1980: The spider fauna of apple trees in central Virginia. Environ. Entomol. 9,247152. -McDawInr, - S. G.; Srnnur.rc, \X/. L., 1,979:Prcdator determination and efficiency on Heliothis oirescens eggs in cotton using r2P. Environ. Entomol. S, 1083-1087. Mrr-rrn, F., 1974: Pavouci fauna repnych poli v okoli Chvalkovic a Nakla na Han6 - Sb. Praci prir. Fak. palack. Univ. Olomouci, 47, Biologica 15,175-1,81,. (Cit. Luczar, J., 1979.) Unpublished results. (Cit. Luczar, J., 1979.) -Mrrrrn, F.; Onnrnr, R., 1975: Soil surface spiders (Araneidae) in a terrestrial reed swamp in southern Moravia (Czechoslovakia). Acta Ent. Bohemoslov. 7 2, 272-285. MIver, S.; KrnrraNr, K.; Sasan.r., T., 1,978: An empirical model of Lycosa-hoppers interaction system in the paddy field. Protection Ecology 1,9-21. Mrvasnrte, K., 1968a: Quantitative feeding biology of Lycosa T-insignita Boes. et Str. (Araneae: Lycosidae). Bull. Nat. Inst. Agric. Sci., Japan, C,22,329-J44. 1968b: Growth and development oI Lycosa T-insignita Boes. et Str. (Araneae: Lycosidae) - under different feeding conditions. Appl. Ent. Zool. J, 81-88. 336 M. Nyffeler and. G. Benz

Mouronn, B. C. ; Rnrcrtr , D. E., 1972: Significance of spider predation in the energy dynamics of forest-floor arthropod communities. Ecol. Monogr. 42, 473498. Mürl-nn, H. J.; BÄnnuaNN, R.; HEINRIcH, Sü.; Menstennn, R.; ScnÄurn, G.; Wrrsacx, $(/., 1978: Zir Strukturanalyse der epigäischen Arthropodenfauna einer Rasen-Katena durch Kescherfänge. Zool. Jb. Syst. 105, 131-1 84. Murraa, M. H., 1973: Comparison of ground-surface spiders in four central Florida ecosystems. Florida Ent. 56, 173-1i6. 1975; in Florida citrus groves. Florida Ent. J8, 83-90. - Spiders MuNrenneN, R. ; CHeoe, H. L., l97Oa: Biological control of the greenbug by the spider Phidippus aud.ax. J. Econ. Ent. 53, 1.712. 7970b: Biology of the crab sprder Misurnenops celer. Ann. Ent. Soc. Am. 63,1718-1722. -Naxauun-a, - M.; Nareuun r , K., 1977 : Population dynamics of the chestnut gall wasp, Dryocos- mus huripbilus Yasumatsu (Hvmenoptära: Cvnipidae). V. Estimation of the effect of predation by Jpid".. on the moitaiity of imaginal'*a'sps based on the precipitin test. Oecälogia, Berl., 27, 97-1.1.6. Nerlsul r. F ., 1976: Factors responsible for the change in the pest status of the tobacco cutworm Spodoptera litura (Lepidoptera: Noctuidae). Physiot. Ecol. Japan 17,527-533. Nnrou, E.,1976: Die Spinnen. In: Einführung in den lntegrierten Pflanzenschutz. Nützlinge in Apfelanlagen. OILB. Internationale Orga-nisation für biologische Bekämpfung schädlicher Tiire und Pflanzen. Sektion Vestpaläarktische Region. Arbeiisgruppe für Integrierten Pflan- zenschutz im Obstbau. Brosch. Nt.3,221-225. Nrcu, A. A.; Hansrav, S. D., 1969: A list of spiders in sugarcane fields in Louisiana' Proc. Louisiana Acad. Sci. 32,50-52. Nrcn, A. A.; Arou-GneDIR, M. F.; SerrrreN, A. G. A., 1975: Zur..Populationsdynamik und Aktivitätszeit der echten Spinnen (Araneida) auf Kleefeldern in Agypten. Anz. Schädlings- kde., Pflanzenschutz, Umwelt schttz 4 8, 1. 61.-1 63. NoHana, K.; Yasuuersu, K., 1965: Observation on the activity of spiders and effect of insecticides on their populations in the citrus groves around Hagi City, Honshu, Japan. Science Bull. Fac. Agric. Kyushu Univ. 23, 1.51.-167. NyrrElrn, M., 1982a: Die ökologische Bedeutung der Spinnen in Forst-Okosystemen, eine Literaturzusammenstellung. [nz. SchadlingsIde., Pflanzenschutz, Umweltschutz SS, 134-] 37 . 1982b: Field studies on the ecological role of the spiders as insect predators in agroecosystems - (abandoned grassland, meadows änd cereai fields). Ph.D. thesis, Swiss Fed. Inst. Technology, Zurich, Switzerland. 1983: Eine Notiz zur ökologischen Bedeutung der Radnetzspinnen als Blattlausprädatoren in - Gärten. Mitt. Schweiz. Ent. Ges. 56,200. NvrrrrEn, M.; Brr.rz, G., 1978: Die Beutespektren der Netzspilnen Argiope bruennichi (Scop.), Araneus quadratus Cl. und Agelena labyrinthica (Cl.) in Odlandwlestn bei Zürich. Revue suisse Zool. 85, 747-757. 1979a Zur ökologischen Bedeutung der Spinnen der Vegetationsschicht von Getreide- und - -Rapsfeldern bei Zürich (Schweiz). Z. rng. Enr. 87,348-376. Nischenüberiappung bezügfich der Raum- und Nahrungskomponenten bei - --1979b:Krabbenspinnen (Araneae: Thomisiilae) und tVolfspinnen (Araneae: Lycosidae) in Mähwiesen. Revue suisse Zool. 86,855-865. 1980a: The role of spiders as insect predators in cereal fields near Zurich (Switzerland). - -Proc. Int. Arachnol. Congr., 8th, Vienna, pp.1.27-131. 1980b: Kleptoparasitiimus von juvenilän Kreuzspinnen und Skorpionsfliegen in den - -Netzen adulter Spinnen. Revue suisse Zool. 87,907-918. 1981a: Freilanduntersuchungen zur Nahrungsökologie der Spinnen: Beobachtungen aus - -der Region.Zürich. Anz. Schädlingskde., Pflanzenschutz, Umweltschutz 54,33-19. 1981b: Ökologische Bedeutung der Spinnen als Insektenprädatoren in Viesen und Ge- - -treidenfeldern. Mitt. dtsch. Ges. allg. angew. Ent. 3,33-35. 1981c: Einige Beobachtungen über die Flucht von adulten Lepidopteren aus den Netzen - -orbiteler Spinnin. Anz. Schäd-iineskde., Pfl anzenschutz, Umweltschutz 5 4, 1 1 3-l 1 4. 7982a: Spinnen als Prädatorerivon landwinschaftlich schädlichen Blattläusen. Anz. Schäd- - -lingskde., Pflanzenschutz, Umweltschutz 5 5, 120-121. Bemerkenswerte Spinnenfunde in \üüiesen und Getreide{eldern bei Zürich. Mitt. - --1982b:Schweiz. Ent. Ges. 5t,97-100. Orurue, C., 1968: Preliminary survey on the spider fauna of the paddy fields in Thailand. Mushi, Fukuoka, 42,89-118. 1973t Spiders in orchards. Shokubutsu-boeki 27,207-21.2. -

I- Spiders in nataral pest control J37

1974: Aeronautic spiders caught by the trap net above paddy fields. Sci. Bull. Fac. Agr., - Kyushu Univ. 29,79-85. 1977 : Studies on the seasonal fluctuation of the spiders in the paddy field ecosystem at Tsuya, - Fukuoka City. Sci. Bull. Fac. Agr., Kyushu Univ. Jl, 133-144. Oruue, C.; 'WöNcsrnI, T., 1973:-Second report on the spider fauna of the paddy fields in Thailand. Mushi, Fukuoka, 47, 1-77. Oruua, C.; Lrn, M. H.; Hoxvo, N., 1978: Fauna of spiders in a paddy field in Suweon, Korea. Esakia .1-1, 81-88. Ornnnc, G., 7960: Spinnen als Feinde der Honigbiene. Imkerfreund 15, 9-1]. PaIr, K. Y.; Krrra, 1. 5., nzl: Survey on the spidir fauna and their seasonal fluctuation in paddy fields of Taegu, Korea. Korean J. Plant Prot. 12, 125-130. Parr, 'SV. H.; Nfu,aroonc, J.; KIu, H. K., 1974 A list of spiders collected in the rice paddy at Milyang. Korean J. PlantProt. 13,24. Parr, V. F.; NeuruNc, J.; PeIx, H. J., 19ß: Spider fauna of the mulberry fields at Suwon, Korea. Korean J. Plant Prot. 12,57-62. Pransn, A., 1946:-Observations on the microfauna o{ the Duke forest. Ecol. Monogr' 16, 127-150. Pncr, S?. B.; 'Vurrcoun, \f. H., 7978: The phenology and populations of ground surface cursorial spiders in a forest and a pasture in t6e south Central United States. Symp. zool. Soc., London, No. 42, l3l-138. Prnrurr.mn, T. R., 1964: Populations of certain insects and spiders on cotton plants following insecticide applications. J. Econ. Ent. 57, 64M44. Prnrrns, E. P.;-SrnnuNG, \(/. L., 1974: Aggregation indices of cotton arthroPods in Texas. Environ. Entomol. J, 598-600. Prr'rExrn, D., 1961: Competition and species-per-genus structure of communities. Ann. Ent' Soc. Am. 54,323-333. Porxrrrc, P. J., 1966: A quantitative field study of predation by the sheet-web spider, Frontinella communis, on Europeän pine shoot moth adults. Canad._J.2oo1.44,265-272. PorrNnc, A., 1968: Un?ersuihungen der terrestrischen Arachnidenfauna im slowenischen Karst- gebirge. Die \üeiden innerhalb der Seslerio auturnnalis-Ostryetum-Assoziation bei Divaca. Biol. Vestn. 16,77-84. PüHnlNcrn, G., 1982: Bedeutung räuberischer Arrhropoden in landwirtschaftlichen Kulturen. (Untersuchungen in unterschiidlich bewirtschafteten Ob-stgärten_ unter besonderer Berück- ii"htig.rtrg dei Spinnen.) Forschungsprojekt-Schlußberichi an das Bundesministerium für Land- und Forstwirtschaft in tü(i ien, 1980 (in prep.). PuruaN, 'W.L.,7967: Prevalence of spiders and iheir importance as predators in Ontario peach orchards. Can. Ent. 99, 1,6V1.70. PurtrnN, 'ü. L.; Hrnrr, D. H. C., 1966: The role of predators and other biotic agents in regulating the population density of phytophagous mites in Ontario peach orchards. Can. Ent. 98,808-821. Ra-arrrarNrN, M.; HuHra, V., 1968: On the spider fauna of Finnish oat fields' Ann. Zool. Fenn' t,254-261. Rrrcrrenr, S. E., 1.974: Thoughts on the ecological significance of spiders. BioScience 24, 352-356. 1984: Spiders as biological control agents. Ann. Rev. Ent. 29, 299-320. -RonrNson,'M. H.; RonrNiox, 8., 1974:\ census of web-building spiders in a coffee plantation at 'Wau, New Guinea, änd an assessment of their insecticidal effect. Trop. Ecol' IJ, 95-107. Rocnns, C. E.; Honqnn, N. V., 1977: Spiders of guar in Texas and Oklahoma. Environ. Entomol. 6,523-524. Roorra, P. M., 1979 Parasites and predators oI Heliothis spp. (Lepidoptera: Noctuidae) in cotton in the Namoi Valley, New Soüth \flales..l. Aust. Entomol. Soc.. 18, 22J-228. RupptnrsnornN, H., 1'96+: Über den Einsariton Decken- und Radnetzspinnen im Forstschutz. Valdhygiene t, 147-153. SaoeNa, G.1.; Keun, M.,7974: Spiders and their prey in citrus orchards. Proc. 61st Indian Sci. Congr. No. 63. SeoeNeiG. L.; S.a.NoHu, D., 1977: Feeding intensity of developmental instars of the spider Marpissa ludbianaensis on the fulgorid pelt of gr_apevines. Sci. and Cult. 43,55V551. SeuNes, G. D.; Renos, R. S., 1975: Imporiance of fiist prey to spiderling survivai in Lycosa sp' (Araneae: Lycosidae). Philipp. Ent. 3,22-29. Serta.tr, P.; MIöne, B.'C., tizi: Spiders: the most effective natural enemies of the brown planthopper in rice. Rice Entomol. Newsl. J, 31. S.q.sÄne, T.iKrnrrerr, K.,7974: Simulations of the population changes of Lycosa in a paddy field. Appl. Ent. Zool. 9, 273-275. 338 M. Nyffeler and G. Benz

Sesen,t, T.; KIntraNt, K.; Kewenena, S., 1970: Assessment of the predatory ability of spiders for comoararive ourDoses. Tap. T. Ent. ZooL 14,144-1.46.

61-64. Sesere, T.; KtntrerI, K.; Unenn, T., 1973b: A preliminary model to simulate the effect of insecticides on a spider-leafhopper system in the-paddy field. Res. Popu! Ecol. I t,9-22' Seurrr, 1., 1936: Inväsion domiciliaire de Rhipicephalus sanguineus et de Teutana triangulosa. Röle ixodiphage des araign6es. Ann. Parasit. Hum. Comp. (Paris) 14, 126-129. Scnrrrr,n, M.', 1974: Experiäentelle Untersuchungen zr.r Bödeututrg der interspezifischen Kon- kurrenz bei 3 \Tolfspinnen-Arten (Araneida: Lycosidae) einer Salzwiese. Zool. lb. Syst. 101, 213-235. ScHerrrn, M.; Hees, L' 1979: Untersuchungen zum Einfluß der Mahd auf die Arthropoden- fauna einer Bergwiese. Drosera (Oldenburg) '79 (1), 1.740. Scnrrrcnn, E. I.; DIrrnrcr, E.J., 1960: Strip-f"t-ittt alfalfa. Calif. Agric. 14,8-9, 15.. Snenu,r., V. K.; Senur, P., 1979; Predatory röle of spiäers in the integrited control of the maize stalk borer, Cbilo partellas (Swinhoe). J. Entomol. Res. J, 299-331 Snrneno, B. M.; Srunr-rNc, \ü. L., 1972:Effecrs of early season applications of insecticides on beneficial insects and spiders in cotton. Tex. Agric. Exp. Stn. Misc.. Publ. No. 1045' SHurov, A., 1938: (Observations on citrus spiders.) (in Hebrew). Hadar, 1l,l-6. SrNcn, G.; Se.woHu, G. S., 1976: New recor-ds of spiders as predators of maize borer and maize jassid. Current Science 45,642. Slrcnr, H. B.; DoNoarB, C. D., 1960: Spider populations in New Jersey apple orchards. J. Econ. Ent.53, 810-814. Srerr,r, P. A.; Crowrn, D. F.; Gnavrs, J. B.; Scnnrrrc, P. E., 1978: Effects of certain herbicides on some insects and spiders found in Louisiana cotton fields. J. Econ. Enl71,477480. SrrvN, J. J., 1959: Use of social spiders against gastrointestinal infections spread by house flies. South African Medical I. 33, 730-711. Susrrrw,rrrw, V.,W., 1939: Untersuchungen über die Bevölkerungsbev/egung des Kiefernspanners (Bapalus(Bunalus piniariasniniarius L.).L-). Mitt.Mitt, Foritwirtsch.For.stwirtsch. u. Forstwissensch. 10.10, 10-51,1'C_51. Suzurr,urr,uKI, Y.;I.; Krnlrer*I,Krnrrer*r,IITRITANI, K., 1.974:l'r/+i ReproductionKeproductron ofOI LycosaLyCOS4 PSeadOAnnAlotApseudoannulata (boesenDerg(Boesenberg andano )[ranq,Strand) (Araneae, Lycosidae) under different feeding condidöns. Jap. J. appl. Ent. ZooL 18,1'66-170. Suzürr,urr. Y.;Y.: Oruue,Oruue. C'C. tgzs:1.975: Spiders inhabitinginhabitine the cabbageTield.cabbaee field. Acta Arachnol. 26,58-43. Te.rsoar, 5., 1.973: Spiders (Araneida) collected in strawberry fields. Norsk. Entomol. Tidsskr. 20,305-307. Tewexe, H., 1973: Ecological studies of spiders in paddy fields in Kinki (I). Studies on the life cycle of spiders, Oedolhorax insecticepi Boes. et-Str. end Enoplognatha japonica Boes. et Str. Acta Arachnol. 2t, lo-1.5. 1975: Ecoloeical studies of spiders in paddv fields in Kinki (ll). Studies on the seasonal - fluctuations "of the two spiders, Oedöthoräx inseaiceps Boej. et Str. and Enoplognatha japonica Boes. et Str. Acta Arachnol. 26,15-57. Ter*exe, H.; Heueuune, T., 1968: Population density of spiders in paddy field during winter. Bull. Coll. Agr. Utsunomiya Univ. 7,73-79.

(Arachnida, Dictynidae) auf Granatapfelbäumen in Agypten. Anz. Schädlingskde., Pflanzen- schutz, Umweltschutz J4, 9G-93. Trneoa, T.,7977: Studies on spider fauna in a tea field (Part 1). Yearround change of numbers of spiders and their name list from Nara Prefecture. Atypus 69, 1'6-20'^ TsR;oe, T.; Yosaluune, K.; IMaNrsHr, M., 1978: Studiei on a spider fauna in a tea field (Part III). Comparative studies on spider fauna of a tea field and the adjacent forests. Atyp'ts 72, 25-30. Trrerrn, K.; SrurNrn, H. M., 1,975: Winteraktive Spinnen auf einem Acker bei Großenzersdorf (Niederösterreich). Anz. Schädlingskde., Pflanzänschutz, Umweltschutz 48-, 184-187. THerrn, K.; AussrnrncHNnn, J.; MuicnNesr, F., 1977: Vergleichende Fallenfänge von Spinnen und Käfern auf Acker- und Grünlandp arzellen bei Innsbruck, Osterreich. Pedobiologia' Jena, 17,389-399. Tr<,rrrn, K.; on Zonoo, I.; Mnvrn, E.; ScHarz, H.; Tnocrn, H., 7978: Arthropoden auf Almflächen im Raum Badsastein (Zenvalalpen, Salzburg, Österreich). Veröff' d. Osterr' Mab-Hochgebirgsprogr. Höhe Tauern. Bd. 2, 196-233.Innsbruck: Univ. Verlag \üagner' TuonNnrrr, R-., 1975: Scorpionflies as kleptoparasites of web-building spiders. Nature 258, 709-711. TxonNnrrr, V. A., 1983 : The distribution and probable importance of linyphiid spiders living on the soil surface of sugar-beet fields. Bull. Brit. Arachnol. Soc. 6,727-136. Spiders in nataral pest control 339

Trscnlen, V., 1958: Svnökologische Untersuchungen an der Fauna der Felder und Feldgehölze (Ein deitrag zur Ökologie äer Kulturlandschaft). Z. Morph. Ökol. Tiere 47, 54-11,4.' 7965: Agrarökologie. Jena: VEB G. Fischer. -Tornrrr, J.-D.; Tavrön, T. H. C; Perrn, R. \(., 1930: The coconut palm moth in Fiji. Imp. Bur. Ent., London (Spiders: p. 153). (Cit. TunNnurr, A. L., 1960.) Tovooe, K. ; YosNrrraune, S., t ggo r A study on the natural enemies of insect pests in paddy field. Kyushu Agric. Res. 28,122-124. TunNnurr, A. L., tgeO: The prey of the spider Linypbia triangularis (Clerck) (Araneidae, Linyphiidae). Can. l. ZooI. 38, 859-873. 1,966: A population of spiders and their potential prey in an overgrazed pasture in eastern - Ontario. Can. I. Zool. 44, 557-583. 7973: Ecology of the true spiders (Araneomorphae). Ann. Rev. Enr. 18,305-348. -Tunrrrsnro, S.-G.; Tooo, J.-!7.; Ceulnnrr, W. Y., 1975: Field activity of selected foliar insecticides against Geocorids, Nabids and spiders on soybeans. J. Georgia Entomol. Soc. .10, 272-277. veN Hoor, R. I., 1971: Energy and nutrient dynamics of spider and orthopteran populations in a grassland ecosystem. Ecol. Monogr. 41, l-26. Vrcirnr'raw, G. i.; Sunderland, K. b., tgzst Arthropods in cereal crops: Nocturnal activity, vertical distribution and aphid predation. I. app. EcoL 12,755-766. Vruesre, A., 1965: Suueaspökti äit'rblik"f"nnäst^kultuurniitudel. 1Über den Sommeraspekt der Spinnenfauna auf den Rulturwiesen.) Eesti NSV Teaduste Akad. Toimetised (Biol.) 14, 329-337. Vn6, J. P., 1953: Unrersuchungen über die ökologische und forstliche Bedeutung der Spinnen im \lalde. Z. ang. Ent. 34. 3"ll-334. \ürrorveun, C.i1 9zS: Über die Bedeutung von Insekten im Ökosystem Laubwald. Mitt. dtsch. Ges. allg. angew. Ent. 1,1.96-204. Vnrrrnn, A. €., tgzl: Studies on the arthropod fauna of alfalfa. V. Spiders (Araneida). Can. Enl 105,425432. 'Wnrrcoun, Y . H., 1974: Natural populations of entomophagous arthropods and their effect on the agroecosystem. In: Summer-Inst. Biol. Control Plint Insects and Diseases. Ed. by Mexvrr-r, F. G.; HannIs, F. A. Jackson: University Press of Mississippi. 150-169. 1975: Ecological role of cursorial ipiders in agroecosystems. Proc. Int. Arachnol. Congr., 6th, - Amsterdam, 1974, p.95. 1980: Sampling Spiders in Soybean Fields. In: Sampling Methods in Soybean Entomology. Ed. - by Koc^lN, M-.; Hrnzoc, D. C. New York-Heidelberg-Berlin: Springer. 544-558. 'Wruicoun, W. H.; Bru, K., 1964: Predaceous insects, spiders and mites of Arkansas cotton fields. Bull. Ark. Univ. 690, 1.-83. '$(inrrcour, tü(i. H.; Taorc, M., 1963: Araneida as predators of the fall webworm. J. Kans. Ent. Soc.36, 186-190. 'Wnrrcoun, \7. H.; ExuNn, H.; HIru, M., 196Ja: Comparison of spider populations of ground stratum in Arkansas pasture and adjacent cultivated field. Proc. Ark. Acad. Sci. 17. 'Wrrrlcoun, V. H.; Exrlrr, H.; HuNrrn, R. C., 1963b: Spiders of the Arkansas cotton {ield. Ann. Ent. Soc. Am. t6,653-660. 'WresueNN, R., 1955: Unteisuchungen an den Prädatoren der Baumwollschadinsekten in Agypten im Jahr 1.951/52. Acta Tropica 12, 222-239. \üolcorr, G. N., 1937: An animal census of two pastures and a meadow in northern New York. Ecol. Monogr. 7, l-90. '\üooos, M. \ü/.; Hannnr, R. C., 1976: Spider populations of a southeast Texas rice field. Southw. Nat. 21,3748. YecrNuua, T., 1965: Spiders found in the paddy field. Plant Protection 19, 361-368. Yeu,r.wo, T., 1977: Sealonal fluctuation of population density of spiders in paddy field in Kyoto Ciry. Acta Arachnol. 27 (special no), 253-260. Yre.nceN, K. Y., 1,975: Prey'aid periodicity of Pardosa ramulosa (McCook) in alfalfa. Environ. Entomol. 4, 137-141.. 1975b: Factors infiuencing the aerial dispersial of spiders (Arachnida, Araneida). J. Kans. Ent. - Soc.48, 403-408. Yrence.N, K. V.; Conrneu, V. R., 1974: An escape barrier for improved suction sampling of Pardosa rarnulosa and Nabis spp. populations in alfalfa. Environ. Entomol. 3, 189-191. Ynnncal, K. V.; Doroarn, C.D.,-ts7+t The spider fauna of alfalfa fields in northern California. Ann. Ent. Soc. Am. 67,681-682. Authors' address: Dr. sc. nat. Menrrx NvrmrBn and Prof. Dr. phil. Gnonc BnNz, Depan- ment of Entomology, Swiss Federal Institute of Technology, ETH-Zentrum, CH-8092 Zurich